What is the sex ratio of harpacticoid copepods in the deep sea?

The sex ratios of deep-sea harpacticoids have been thought to be greatly skewed toward females. The representation of males is notably more equitable (approximately 1:2) at three deep-sea sites (San Diego Trough: 32°52.4N, 117°45.5W, 1 050 m, January 1987; Porcupine Seabight: 51°36.85N, 12°57.30W, 1 369 m, August 1984; northwest Atlantic, 40°27N, 62°20W, 4 820 m, July 1982, June 1983). This ratio conceals extensive variation at the species level. For some species, males are unknown. For other species, males are more numerous than females. Sampling bias against males can occur and may explain the more extreme reports of male rareness in the literature.     

Food threshold territoriality in a water strider (Gerris remigis)

Summary Summer generation 3rd, 4th and 5th instar nymphs plus adults of Gerris remigis were satiated for 2 days in a laboratory tray then deprived of food. Within 1/2–2 days, 19 of 27 nymphs and 10 of 30 adults began to exhibit territoriality, continued being so for 3–9 days, then ceased shortly before becoming quiescent. In the field, muscid flies fed to different territorial striders at a rate of 0, 1, 2, or 3 flies/day, resulted in 8 of 10 striders (at 2 flies/day) and 10 of 10 striders (at 3 flies/day) ceasing territoriality, whereas 5 of 8 controls (at 0 flies/day) remained territorial. Thus, lower and upper food thresholds were demonstrated, the upper threshold approached both from above (laboratory study) and below (field study).     

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.     

Baboon loud calls advertise male quality: acoustic features and their relation to rank,age, and exhaustion

Free-ranging adult male baboons give loud two-syllable wahoo calls during dawn choruses, interactions between groups, when chasing females, and in aggressive interactions with other males. Previous research has shown that the rate and duration of these contest wahoos are correlated with a males competitive ability: high-ranking males call more often, call at faster rates, and call for longer bouts than do low-ranking males. Here we report that acoustic features of wahoos also reveal information about male competitive ability. High-ranking males give wahoos with higher fundamental frequencies (F0) and longer hoo syllables. Within-subject analyses revealed that, as males fall in rank, the hoo syllables tend to shorten within a period of months. As males age and continue to fall in rank, F0 declines, hoo syllables shorten, and formant dispersion decreases. Independent of age and rank, within bouts of calling F0 declines and hoo syllables become shorter. Because wahoos are often given while males are running or leaping through trees, variation in these acoustic features may function as an indicator of a males stamina. The acoustic features of contest wahoos thus potentially allow listeners to assess a males competitive ability.Communicated by C. Nunn     

Genetic evidence that the “Padstow mussel” is Mytilus galloprovincialis

Electrophoretic analysis of loci controlling a variety of enzymes has been applied to samples of the Padstow mussel and typical Mytilus edulis L. living strictly sympatrically at Rock, Cornwall, England, in order to resolve the disputed status of the Padstow mussel. Small samples indicated similar monomorphic states at the GPDH, TO, MDH-2 loci and weak polymorphism at the 6 PGD locus in both types of mussel. The MDH-1 locus may be weakly polymorphic in the Padstow mussel and monomorphic in M. edulis. Large samples assayed for AP, LAP, PGM, and PHI produced data showing very large and highly significant differences in allele frequencies at three of these loci between the two groups of mussels separated on anatomical characters. At the LAP locus, significant deficiencies of heterozygotes were observed in both groups of mussels. A small percentage of mussels from Rock are difficult to assign with certainty to one or other group on anatomical and morphological criteria, but the genetic evidence indicates that most, if not all, such specimens are M. edulis. These genetic differences make it highly improbable that any significant degree of genetic exchange occurs between the two groups in nature and, taken together with the evidence of genetic resemblance of the Padstow mussel to M. galloprovincialis from Venice and that of Rock M. edulis to M. edulis, from the Gower peninsula of Wales we conclude that the Padstow mussel is indeed M. galloprovincialis.     

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     

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.     

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.     

Alternative mating tactics and the law of Diminishing Returns: The Satellite Threshold Model

Summary The mating systems of many species comprise at least 2 alternative male mating tactics — broadly termed Dominant and Satellite tactics here. But, there exists no satisfactory conceptual framework in which to explain their co-occurrence and persistence.A Dominant male's ability to copulate is a positive, but decelerating, function of either his territory's attractiveness or his own attractiveness to females. This Diminishing Returns Effect affords mating opportunities for Satellite males associated with the most attractive sites (or males). A graphical Satellite Threshold Model, analogous to the Polygyny Threshold Model (Orians 1969), is developed to predict quantitatively the presence or absence of Satellite male tactics.Five predictions of the Model are supported quantitatively by data from dragonfly and anuran social systems and, more generally, by evidence from a wide variety of species.The Model should apply both to circumstances in which males are able to alternate between tactics and the instances in which males make an irrevocable tactical decision.     

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.     

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.     

Sexual conflict in the house sparrow: interference between polygynously mated females versus asymmetric male investment

Summary The reproductive success of female house sparrows mated with polygynous males depended to some extent on the aid received from their mates. Polygynous males fed nestlings at the same rate as monogamous males although polygynous males gave aid almost exclusively to one of their mates (the preferred). As a consequence, the number and quality of young raised by preferred females were similar to those of monogamous females, whereas nonaided females experienced a reduction of clutch size, hatching success, and fledgling quality. Males aided the harem female that hatched her clutch earliest (in 11 out of 12 cases). However, harem females that laid eggs earliest (i.e., the potential preferred) lost eggs and/or nestlings more frequently than harem females that delayed laying. This resulted from competing females' infanticide suited for redirecting males' aid. All polygynously mated females delayed the start of a new clutch and reproduced fewer times per season than monogamous females. Consequently, monogamy seems to be the optimum mating situation for females; whereas polygynous males raised more young per year than monogamous males, interference between harem females reduced their maximum expected success.     

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.     

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     

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     

A mechanism for female choice of large males in the treefrog Hyla chrysoscelis

Summary In this study, we combine both field and laboratory experiments to address the effects of female preference for certain call characteristics on a large-male mating advantage in the treefrog H. chrysoscelis. In laboratory-choice experiments, females always chose the call with the lower fundamental frequency when call rate and call intensity were held constant and the difference in frequency between the two calls was 15%. The lower frequency call was preferred by 8 out of 12 females when the difference in fundamental frequency was 7.4%. These results are consistent with field comparisons of the size of unmated males calling within 2 m of a mated male: Male body size was negatively correlated with fundamental frequency and the greater the size difference, the more likely that the larger male mated. In field choice experiments, females preferred males with higher call rates. Since size differences between males used in this experiment averaged only 2.3 mm, we would not expect the fundamental frequency of a male's call to be the best predictor of mating success. Laboratory results demonstrated that call rate could override female preference for the low frequency call over the high frequency call, while intensity could at least dilute this preference. However, individual males in the field varied both call rate and the call intensity as perceived by the female. We suggest that the interaction between call rate, male size and mating success should be studied further through the use of field-choice experiments.     

Eusociality and extraordinary sex ratios in the spider Anelosimus eximius (Araneae: Theridiidae)

Summary Colonies of Anelosimus eximius in Panama had an average sex ratio of 0.15±sd 0.09, i.e. about five females for each male. The sex ratio in egg sacs reared was even lower (0.08±0.01), as was that of immatures in newly founded colonies (0.12±0.05). The possible mechanisms responsible are discussed. Mature colonies had an average ratio of 17 females and 2 males for each egg sac present (range: 2–91 females, 0.2–8.2 males) and contained a large proportion of females which were not inseminated but which presumably help. Since both sexes are diploid, arrhenotoky can be ruled out and it is assumed that some females do not come to reproduction, the proportion depending on the availability of resources. This mechanism may enable entire colonies to survive lean times.     

“Alternative” mating behavior in the swordtails Xiphophorus nigrensis and Xiphophorus pygmaeus (Pisces: Poeciliidae)

Summary The swordtail Xiphophorus nigrensis exhibits three relatively discrete male body-size classes that derive from allelic variation at the Y-linked pituitary (P) locus. Previous studies have shown that larger males have greater relative reproductive success, and that females prefer large males. We describe the mating behavior utilized by males of each size class during individual encounters with females. Small males rely on chase behavior, similar to the alternative mating behavior classified as sneaker in small males of other species. Large males court and intermediate-sized males court or chase, depending on their body size. There is a strong correlation between P alleles for small size (s) and large size (L) with chasing and courting, respectively. The relationship between mating behavior of males of the genotype I is ambiguous. In the closely related species X. pygmaeus, males are of size similar to smaller X. nigrensis males. Paradoxically, these males do not show the courtship display that typifies larger male X. nigrensis and many other species of swordtails, but instead often employ chase behavior identical to the alternative mating behavior in small male X. nigrensis. We suggest that historical and genetic constraints, in addition to current selection forces, might be important factors in explaining the existence of alternative mating behavior in X. pygmaeus.     

Sex ratio,monogamy and breeding success in the Midas cichlid (Cichlasoma citrinellum)

Summary The Midas cichlid (Cichlasoma citrinellum) is a large, aggressive monogamous fish from the Great Lakes of Nicaragua. It lives in an ecosystem where breeding sites are at a premium and where breeding success can be low. I tested the strength of the pair bond by manipulating the sex ratios in captive groups, using ratios of 1:1, 2:1, 4:1, 1:2 and 1:4 (:). No polygamy was seen at any of the ratios. Departure from a ratio of 1:1 decreased breeding success for those pairs that formed. Both sexes appeared to enforce the pair bond but did so in different ways: females attacked males and females equally but males attacked other males significantly more than they did females. These results, coupled with field observations, indicate the Midas cichlid is obligated to monogamy by the demands of competition for breeding sites, the need for two parents to protect the fry and by energetic constraints.     

Social control of deception among status signalling house sparrows Passer domesticus

Summary Motivation to fight and willingness to risk the use of dangerous weapons, two factors which can be decisive in predicting the winners of asymmetric animal contests, are readily bluffed in display. When contests are decided by motivation to fight or daring, more reliable information about contest asymmetries may be gained by watching actual fights than by watching displays. By presenting mounts to territorial red-winged blackbirds and scoring how aggressively a territory holder attacks the mount, then recording any subsequent intrusions over established boundaries, I showed that neighbors are likely to intrude upon focal males that fail to attack the mount vigorously. This result suggests that red-winged blackbirds watch contests to assess the RHP of neighbors.