The role of male vs male interactions in maintaining population dialect structure

Summary Nonmigratory populations of Whitecrowned Sparrows in coastal California exhibit dialects in territorial male songs that are stable in space and time. By field playback experiments, we tested a prediction from the hypothesis that male aggressive interactions prohibit mixing of song dialects. Playback of the home dialect Clear song to territorial males singing the Clear dialect resulted in less response than that given to playback of the neighboring Buzzy dialect. Response to both Buzzy and Clear dialect songs by target Clear males, however, was greater than that given to the Bodega dialect recorded 55 km away (Figs. 2 and 3). We conclude that these results are consistent with the hypothesis that male-male aggressive interactions play an important role in reducing dialect mixing.     

The evolution of cheating and selfish behavior

Summary We examine the necessary conditions for the spread of genes that determine selfish and cheating behaviors and the rate of spread of these genes through structured populations, in order to address the question of the invadability of altruistic systems by anti-social mutations. We find that, although cheaters always have a higher relative fitness than altruists within groups, population structures which permit the evolution of altruism also preclude invasion by anti-social mutations. These results are related to a discussion by Hamilton (1971) concerning the limits to the evolution of altruistic and selfish behaviors.     

Testing the ‘agonistic buffering’ hypothesis

Summary Data are presented from a 15-month study on triadic male-infant interactions (agonistic buffering) among wild Barbary macaques, and the agonistic buffering hypothesis reevaluated. The sociometrics of triadic interactions derived from the distribution of 535 interactions among individually known adult and subadult males showed that there were significant individual male differences in the frequency of initating and/or receiving triadic interactions, but there were no such differences between the high and low ranking male subgroups (Tables 2–4).Males did not choose each other equally for triadic encounters. Each male had a different set of three other males (out of 11 possible) that he preferred to approach for a triadic encounter; and each male received triadic approaches essentially from only three males (Tables 4 and 5). Each male showed a specific preference for which infant to select for participation in the triad, and in fact, males who preferred one another for triads also preferred to use the same infant in these encounters. These males that preferred each other and the same infant both had a special care-taking relationship with that preferred infant (Tables 6–7).Possessing an infant per se seemed to be irrelevant in whether a male would approach or be approached for a triad. In 78% of all triads, males separated immediately after termination of the triad. When the agonistic buffering hypothesis is reexamined against the data in this study, it appears that it cannot adequately accommodate the thesis that it serves to regulate dominant/subordinate relations among males. Rather males choose to participate with each other in agonistic buffering because, and by means of, a shared, common, and special care-taking relationship with the same infant. The indiscriminate use of the terminology agonistic buffering to describe multiple-male/infant interactions in this species should be dropped.     

Are communal pukeko caught in the prisoner's dilemma?

Summary The prisoner's dilemma of game theory provides a possible explanation for communal breeding in the pukeko (Porphyrio porphyrio, Aves) where group breeding birds are at a reproductive disavantag compared to pairs. Territorial defence is largely the realm of males. A male who defect on neighbours by taking in an extra male defender and so becoming communal stands to gain territorial area and possibly higher reproductive success. Neighbours are forced to follow suit if additional males are available or lose their territory and therefore their breeding opportunities. If neighbours also admit an extra defender, the initial defector loses his advantages and all males end up with the lower breeding success of communal groups. Such defection to a communal habit appears to be the only stable solution.     

Lead-based paint in dwellings: The potential for contamination of the home environment during renovation

The quantity and particle size characteristics of lead in dust released during three different paint removal techniques was determined under controlled conditions and in situ in a dwelling. Air-lead and deposited dust-lead levels were highest after sanding but burning-off and hot-air removal methods also produced significant contamination. The importance of dust particle-size and lead is discussed in relation to the potential hazard to home renovators via inhalation and to children via the hand-to-mouth route.This paper is also being published in Proceedings of the Conference on Lead in the Home Environment and is one of several selected from SEGH sponsored conferences for simultaneous publication in this journal.     

The erect ‘penis’ is a flag of submission in a female-dominated society: greetings in Serengeti spotted hyenas

In East Africa, spotted hyenas live in large clans in a highly structured society dominated by females. A clan is a fission-fusion society where members are often solitary or in small groups. Spotted hyenas have a ritualized greeting during which two individuals stand parallel and face in opposite directions. Both individuals usually lift their hind leg and sniff or lick the anogenital region of the other. The unique aspect of greetings between individuals is the prominent role of the erect penis in animals of both sex. Female spotted hyenas have fused outer labiae and a pseudo-penis formed by the clitoris which closely resembles the male penis and can be erected. During greetings subordinates signalled submission with gestures which were not necessarily reciprocated by the dominant participant. Asymmetries were most pronounced in greetings between adult females where the probability of asymmetries increased with the divergence in rank between partners. Greetings between adult females and males were uncommon and restricted to males above median rank, principally the alpha male. Models of primate affiliative behavior assume that benefits derived from social relationships with different individuals are not equal and that individuals are selected to maximize the benefits they receive from social relationships with others. The observed distribution of greetings between partners of different rank matched the predictions of these models. An examination of non-adaptive hypotheses on the evolution of the pseudo-penis demonstrated that the conventional scenario linking (initial) virilization of female genitalia with selection for female dominance does not explain either the initial virilization, nor the evolution of the pseudo-penis to its current form and use. We sketch a new scenario that links (1) initial virilization to the occurrence of neonatal siblicide amongst members of a twin litter, and (2) costs of maintenance, pseudo-penile control over copulation and male submission. Our analysis confirms previous adaptive hypotheses on the function of the pseudo-penis in greetings and suggests new hypotheses to account for hitherto unexplained features. Correspondence to: M.L. East     

Trunk trails and the searching strategy of a leaf-cutter ant,Atta colombica

Summary Atta colombica uses chemical mass recruitment that allows the rapid exploitation of resources. Most foragers thus search only within patches. Accumulation of extra foragers at patches results in sampling of alternate food items and area-restricted search as patch resources are depleted.Individual workers have a higher probability of removing a leaf fragment the earlier they arrive at a bait. Workers that arrive when much of the resource is gone travel further on the bait (within the patch) but do not spend significantly more time at the patch. They give up after 50–80s.Foraging effort is centered on the extensive trail system, not on the nest a predicted by time and energy foraging models. Search effort is also trail centered. The probability that an item will be discovered decreases with distance from the trail and increasing litter depth. Trail traffic and trail quality together mave no significant effect although this may be because they act antagonistically.Economic considerations predict that trials should be built to high quality and very productive sites. If trails are built as a result of recruitment and recruitment reflects patch quality and productivity, characteristics of forage sites are physically embodied in the trail system.Leaf cutter foraging is better understood as a long term optimization that effectively exploits resources over the lifetime of the colony than as prudent predation that husbands resources.     

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     

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     

On the evolution of male workers in the Hymenoptera

Summary The effects of brood sex-ratio (investment ratio) and the presence of laying workers on relatedness in the Hymenoptera are analyzed. It is shown that the conditions of brood composition that generate degrees of relatedness favorable to the evolution of eusocial type helping behavior among females will select against such helping behavior among males, and vice versa (see Fig. 1). Thus, societies in the Hymenoptera can be expected to have male workers or female workers, but not both. It is argued that the conditions leading to degrees of relatedness favorable to male helping are quite restrictive and unlikely to be met in haplodiploid species.The presence of laying workers is shown to lead to biasses in relatedness such that females may be selected to be workers even when the sex-ratio is male-biassed. This result sheds new light on a possible pathway to eusociality in the Hymenoptera. It is argued that offspring parasitism of the natal nest may have been important in the evolution of eusociality.     

Comparisons of the feeding activity and diets of male and female cunners Tautogolabrus adspersus (Pisces: Labridae)

The diel distribution of feeding activity during the spawning season was compared for territorial male and female cunners Tautogolabrus adspersus in Conception Bay, Newfoundland, Canada between June and August, 1978. Territorial males feed significantly less often than females and concentrate their feeding activities in the morning, whereas females feed as frequently in the afternoon as in the morning. Based on the contents of alimentary tracts, territorial males, non-territorial males, and females have different diets. Since these three groups share the same habitat, the observed dietary differences probably reflect differences in their foraging behaviour. The relevance of these findings to the concept of time minimizer and energy maximizer is discussed.     

Song as an attractant for male and female European starlings,and the influence of song complexity on their response

Summary We tested the hypothesis that song attracts females and repels males in the European starling. We broadcast recorded song from speakers attached to nestboxes, while paired boxes with silent speakers served as controls. As predicted, females were attracted to the song boxes. Contrary to prediction, males were also attracted to the `song' boxes. Singing by male starlings may be costly because it attracts competitors for limited nesting sites, but the cost cannot be avoided due to the need to attract a mate. In a second experiment simple song (composed of 20 different phrase types) and complex song (40 phrase types) were played simultaneously. More males were captured at boxes where simple song was played. Song complexity may function as an indicator of male quality and be used by male starlings to assess potential competitors.     

The use of ICP-MS for the analysis of natural waters and an evaluation of sampling techniques

Analytical and sampling methods for a reconnaissance survey of river waters in former metal mining areas of Wales, UK, are considered. A semi-quantitative mode of analysis with a single internal standard, possible by inductively coupled plasma mass spectrometry (ICP-MS) is compared with more conventional quantitative ICP-MS methods using external calibration from aqueous standards. Semi-quantitative analysis can be much quicker than a more rigorous quantitative approach and gives values which agree with fully quantitative data to within 30%, indeed, agreement is often much better than this. A comparison of samples filtered and acidified in the field with untreated samples is also presented. This shows that in most cases it is unnecessary to filter and acidify samples if the storage time before analysis is short (in this case typically less than four days).     

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     

Importance of the fatty acids 20:5ω 3 and 22:6ω 3 in the diet of plaice (Pleuronectes platessa) larvae

The requirement of plaice (Pleuronectes platessa L.) larvae for the fatty acids 20:5 3 and 22:6 3 was studied. Larvae were reared from first feeding to beyond metamorphosis on Artemia sp. nauplii (EG brand), whose nutritional content had been manipulated by enrichment. Some larvae were fed Artemia rich in 20:5 3 and 22:6 3, and others were fed nauplii with no 22:6 3 and low levels of 20:5 3 (1.9% of the total fatty acids). The differences in diet had no significant difference on growth or survival of the plaice larvae. The results indicate that the plaice larvae in this study did not require 22:6 3 in their diet, even when the levels of 20:5 3 were low.     

Optimal foraging and territorial defence in the Great Tit (Parus major)

Summary We describe an experiment designed to investigate the trade-off between foraging and territorial vigilance in the Great Tit. Captive territorial male Great Tits were observed while foraging in a large indoor aviary. They obtained food from two operant patches in which the supply of food was gradually depleted during a visit. We predicted that during control sessions the birds would switch between patches in such a way as to maximise their overall feeding rate. In experimental sessions, we introduced briefly a rival male as an intruder at the start of the test. The foraging male could see the rival only when travelling between patches and not while feeding within patches. We predicted that during experimental sessions birds would switch between patches more often than in control tests, sacrificing food intake for territorial vigilance. Three of the four males tested behaved in approximately the predicted manner. We discuss the use of an inverse optimality argument to provide a calibration of feeding against the benefit resulting from territorial vigilance.     

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     

Discrimination between aluminium held within vegetation and that contributed by soil contamination using a combination of Electron Probe Micro Analysis (EPMA) and Inductively Coupled Plasma-Atomic Emission Spectrometry (ICP-AES)

Aluminium concentrations in NIST (formally NBS) Standard Reference Material Tomato Leaves and Citrus Leaves were determined by ICP-AES after acid digestion. The normal total acid attack (nitric and perchloric acids) gave very low recovery (40%) of Al in NIST tomato leaves, but not for citrus leaves. The contribution of Al from Al-rich soil particles in both tomato leaves and citrus leaves was estimated semi-quantitatively by computer-controlled EPMA. The Al held in these particles corresponded in approximate concentration to the shortfall between the acid soluble component determined in this study and the certified value. Analysis by EPMA can provide, therefore, a method of estimating Al contributed by soil contamination to plant materials. The limitations of the NIST tomato and citrus leaf reference material for the validation of methods for the determination of Al in vegetation are demonstrated.     

Stability of eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) depth limits: influence of habitat type

Seagrass meadows are generally considered stable although few studies have specified and tested this statement. On the basis of a large monitoring dataset from Danish coastal waters, we aimed to test whether the stability of deep eelgrass populations changes along a eutrophication gradient extending from inner bays over outer bays to open coastal waters. We defined stability in terms of the stability properties of constancy, resilience, and/or persistence. Our data allowed us to investigate the stability property constancy expressed as temporal variability in eelgrass depth limits over a time scale of 10 years. We hypothesised that colonisation depths were large and relatively constant along open coasts because of low and relatively constant levels of nutrients and turbidity. Conversely, colonisation depths were hypothesised to be low and variable in protected bays due to higher and more variable levels of nutrients and turbidity. We found that depth limits increased from inner bays towards open coastal waters, matching declines in nutrient concentration and increases in water clarity and oxygen concentration. Stability expressed as constancy of depth limits did not differ significantly between habitat types, and neither did stability of physicochemical variables. However, when data from all habitat types were analysed together, they showed that eelgrass populations at the depth limit were significantly more constant and thus, in this respect, more stable when occurring in deep waters as compared to shallow waters. Areas of good water quality may thus obtain the double benefit of deeper-growing and more stable eelgrass populations. The most likely reason why this pattern did not appear at habitat-type level is that the habitat types studied represented wide spatial variation in water quality and depth limit. We conclude that the question of whether eelgrass populations are stable depends on the stability property and the ecological situation in question. Populations may be considered unstable in terms of inter-annual variation but stable in terms of long-term persistence. Therefore, the common statement that eelgrass populations are stable is not universally true.Communicated by M. Kühl, Helsingør     

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