The effects of castration, sex ratio and population density on social segregation and habitat use in Soay sheep

We analysed 16 years of census data gathered on the island of Hirta (archipelago of St. Kilda) to investigate the effects of castration, population density, sex ratio, season and group type on habitat use and social segregation of Soay sheep. From 1978 to 1980, 72 male lambs were castrated. We used this experiment to study how a change in reproductive status could affect sociality and habitat choice of these males. Males, females and castrates were all segregated outside the rutting season in autumn. Castrates were the least segregated from females in spring and summer but were most segregated from them during the pre-rut. The more equal the sex ratios, the higher was the degree of social segregation. The three sex classes used similar habitat types, namely, Holcus agrostis, Agrostis festuca and Calluna habitats. Holcus agrostis and Agrostis festuca were top- and second-ranked in female and castrate habitat use, while Holcus agrostis and Calluna were the two top habitat types used by rams. It is unclear why males included Calluna heath habitats, but it cannot be excluded that they might have shifted their use depending on forage availability. A lack in differences in habitat use between castrates and females suggests that body size differences alone cannot be the driving factor for habitat segregation in male and female Soay sheep and that there are reasons other than body size that could motivate reproductive males to use additional habitat types, such as Calluna heath. Although habitat use shifted from one habitat type to the next between low- and high-population-density years and between seasons, there was no clear link between population density and how different groups (male, female or castrate) used these areas. We discuss effects of reproductive status, population density and sex ratio on social segregation and habitat use and suggest that these factors need to be taken into account when investigating causes of sexual segregation in ungulates.     

Muscle esterase variation and size variation in the sand launce Ammodytes dubius

Esterase variation detected in homogenates of muscle tissue taken from the sand launce Ammodytes dubius in April, 1970 and 1971 is described. Eight alleles were found segregating at a single locus, and the frequencies of these alleles were homogenous in population samples taken on the Emerald Bank, off Nova Scotia, Canada, in successive years. When ordered by their electrophoretic mobility, the profile of the frequencies of these alleles was unimodal and symmetric. In both of the population samples, there were significant excesses of homozygotes and deficencies of heterozygotes. There was an association between genotype and size of individuals among homozygous genotypes; individual homozygous for the most common, intermediately-migrating allele were larger than homozygotes bearing alleles with relatively fast or slow mobilities.     

Sex-biased maternal expenditure in Rocky Mountain bighorn sheep

Summary We examined maternal energy expenditure in Rocky Mountain bighorn sheep to test the hypothesis (Trivers and Willard 1973) that individual sons in polygynous mammals should obtain a larger maternal subsidy. In accord with theory, males weighed more at birth (an index of relative prenatal expenditure), tended to suckle more between 40 and 100 days of age (an index of relative postnatal expenditure) and imposed greater delays on their mothers' return to estrus in the subsequent breeding season (an index of relative total preweaning expenditure). Mothers rather than offspring appeared to have primary control over postnatal expenditure. The effect of maternal reproductive effort on return to estrus was cumulative over a period of 3–5 years so that ewes making relatively large expenditures gave birth progressively later. Ewe and lamb mortality was associated with late conception in the previous rut. Thus, differential reproductive effort by ewes in this population appeared to entail a fitness cost involving reductions in both offspring and maternal survivorship. Correspondence to: J.T. Hogg at the present address     

Birth sex ratios in sheep over six lambing seasons

Summary The birth sex ratio of a commercial flock of Suffolk × sheep, Ovis aries, were studied over six consecutive lambing seasons. In all, data from 1820 lambs were recorded and analysed. The overall birth sex ratio was 49.56% male lambs. Significantly more males (56.23%) were born to ewes with single lambs than to ewes with like-sex twins (47.73%). Significantly more female lambs were born to ewes with triplet births (41.79% male). A significant difference in the birth sex ratio was observed between the first and second half of the lambing season, among like-sex pairs of twin lambs. More males were born in the first half (51.43%) and significantly more females (43.59% male) in the second half of the lambing season. The results are discussed with reference to the theory of Trivers and Willard (1973).     

Geographic variation in resource dominance-discovery in Brazilian ant communities

A predictive framework for the ecology of species invasions requires that we learn what limits successful invaders in their native range. The red imported fire ant (Solenopsis invicta) is invasive in the United States, Puerto Rico, Australia, New Zealand, and China. Solenopsis invicta appears to be a superior competitor in its introduced range, where it can cause the local extirpation of native species, but little is known about its competitive ability in its native range in South America. Here we examine the competitive ability of S. invicta for food resources in three widely separated Brazilian ant communities. Each of these communities contains 20-40 ant species, 8-10 of which were common and frequently interacted with S. invicta. S. invicta at all three sites was attacked by several species-specific phorid parasitoids, and at one site, two other species were attacked by their own specialized parasitoids. We examined interactions in these local communities for evidence that trade-offs among ant species between resource dominance and resource discovery, and between resource dominance and parasitoid vulnerability facilitate local coexistence. The trade-off between resource dominance and resource discovery was strong and significant only at Santa Genebra, where parasitoids had no effect on the outcome of confrontations at resources. At Bonito, parasitoids significantly reduced the ability of S. invicta, which was the top-ranked behavioral dominant, from defending and usurping food resources from subordinate species. In the Pantanal, S. invicta ranked behind three other ant species in a linear hierarchy of behavioral dominance, and lost the majority of its interactions with a fourth more subordinate species, Paratrechina fulva, another invasive species. Parasitoids of S. invicta were uncommon in the Pantanal, and did not affect its low position in the hierarchy relative to the other two sites. Parasitoids, however, did affect the ability of Linepithema angulatum, the top-ranked behavioral dominant in this community, from defending and usurping resources from behavioral subordinates. These results indicate that both interspecific competition and trait-mediated indirect effects of phorid parasitoids affect the ecological success of the red imported fire ant in its native range, but that the relative importance of these factors varies geographically.     

Copulatory tactics in relation to sperm competition in Rocky Mountain bighorn sheep

Summary Sperm competition arises in the bighorn mating system because receptive ewes are some-times defended and mated in sequence by a series of increasingly dominant rams and because subordinate (coursing) rams commonly succeed in forcing matings with defended ewes. To identify potential components in a male strategy of sperm competition, the social circumstances and schedule of copulation among free-ranging bighorns were investigated. Three such components emerged: (i) Defending and coursing rams copulated with estrous ewes at extraordinarily high rates, presumably to increase the proportion of their own versus rivals' sperm present at fertilization by simple numerical preponderance at insemination. (ii) Coursing but not defending rams concentrated copulations later in the period of estrus (sensu this study). Evidence from domestic sheep suggests that this may correspond to the period of maximum female fertility. A greater risk and/or reduced benefit associated with forced versus cooperative copulation may force coursing rams to restrict copulation to periods of highest fertility, whereas defending rams may gain even from copulations outside this period. (iii) Defending rams copulated at unusually high rates immediately after bouts of successful coursing. Such retaliatory copulations were associated with reduced amounts of courtship and were more likely to occur if the defending ram had copulated infrequently recently. The number of retaliatory copulations was independent of the number of preceding forced copulations, probably because defending rams rationed ejaculates in anticipation of future coursing. The function of retaliation is not clear, although the literature on domestic sheep suggests the possibility of some form of mechanical displacement of rival sperm. Unlike females of many ungulate species, bighorn ewes were typically receptive to cooperative copulation over a period of days. Although persistent courtship by defending rams may coerce such extended receptivity, it is also possible that ewes, facing unavoidable forced matings by coursing rams, gain genotypic benefits for offspring by providing prolonged copulatory access to relatively dominant defending rams.     

Evolution of variation in male secondary sexual characteristics

Summary The male pied flycatcher Ficedula hypoleuca exhibits sexual dimorphism in its plumage colour, varying from a female-like brown to jet black. The evoltution of this variation in male plumage colour can be explained by at least eight hypotheses viz., (I) neutral mutation; (II) individual recognition; (III) three forms of inter-sexual selection; (IIIa) mate selection for phenotypes, (IIIb) Fisherian selection, (IIIc) handicap selection; (IV) intrasexual selection; (V) delayed maturation; and (VI) female mimicry. The assumptions and predictions derived from all these hypotheses were tested by analysing the observed variation in male plumage colour in relation to age, body size, physical condition, survival rate, aggressivity in territorial defence, territorial quality, female choice of mate, sex ratio, and reproductive success. We found that: males became blacker with age; black males were larger than browner ones; however, browner males survived better between breeding seasons than did blacker ones; black males were more aggressive against black than brown intruders; blacker males occupied better territories; blacker males were paired earlier than browner males; and finally blacker males produced heavier offspring than did brown males. A reasonable conclusion from the results of these analyses is that four of the hypotheses considered, viz. mate selection for phenotypes (IIIa), Fisherian selection (IIIb), handicap selection (IIIc), and intra-sexual selection (IV) could not be rejected. We therefore suggest that the evolution of the observed variation in the secondary sexual characteristics of the male pied flycatcher can be explained by a combination of these hypotheses.     

Genetic variation in the mangrove periwinkle Littorina angulifera

Twenty populatios of Littorina angulifera, inhabiting islands composed of the mangrove tree Rhizophora mangle, were assayed at an esterase locus to determine whether genetic differentiation was associated with distance between populations. It was predicted, on the basis of larval dispersal in this species, that genetic differentiation between populations on islands separated by long distances should be greater than those on islands located near each other. A chi0square test of homogeneity revealed significant differences in esterase gene frequencies among the 20 island populations. However, there was no association of distance between islands and genetic heterogeneity. In addition, a cline in gene frequency was found to be assiciated with latitude. Factors responsible for the observed pattern of heterogeneity at the esterase locus are discussed.     

Individual variation in winter foraging of black-capped chickadees

Summary Wintering black-capped chickadees (Paridae: Parus atricapillus) in northwestern Massachusetts showed a high degree of individual variation in foraging behavior. After accounting for the effects of different habitats and weather conditions, individual differences comprised 6–17% of the total observed variation in four measures of foraging location and rate of feeding. Differences between age and sex groups were not significant and explained comparatively little variation (0.0–1.4%). The chickadees did not fall into a few distinct behavioral categories but instead showed continuous variation on all measures of foraging behavior. It appeared that some variation among individuals was a consequence of behavioral convergence within social groups, since birds that were observed together were more similar in their foraging than expected by chance, after taking habitat differences into account. Our results therefore do not support the interpretation that individual variation in feeding behavior serves to reduce exploitation competition within social groups.     

Genetic variation in the eel: a critique

A number of papers have dealt with protein variation in samples of eels of the genus Anguilla from throughout the North Atlantic Ocean. Examination of these published data from the standpoint of observed electrophoretic variations as well as statistical comparisons with genetic models, suggests that most genetic inferences concerning eels have been unjustified.     

Family structure and variation in reproductive success in blackbirds

In avian families, some offspring are rendered unequal by parental fiat. By imposing phenotypic handicaps (e.g., via asynchronous hatching) upon certain of their offspring and not others, parents structure the sibship into castes of advantaged “core” offspring and disadvantaged “marginal” offspring that results in an asymmetric sibling rivalry. Here, I show how this family structure scales up to population level reproductive consequences. In a 17-year study of red-winged blackbirds (Agelaius phoeniceus), I show that year-to-year variation in the number of surviving offspring is driven primarily by variation in the number of marginal offspring at hatching and their posthatching survival. Clutch size, core brood at hatching, and fledging varied little from year to year and had little direct effect on year-to-year variation in total brood size at fledging; conversely, variation in the size of the marginal brood at hatching and at fledging was much greater. Marginal but not core brood size at hatching rose with mean clutch size; in years where parents laid larger average clutches they did so by adding marginal progeny. The mean posthatching survival of marginal offspring was always lower than that of core offspring in a given year, and there was no overlap in the distributions. The highest mean survival of marginal offspring across years fell below the lowest mean survival of core offspring; broods were deeply structured. There was an overall female bias among fledglings, and the sex ratio varied across years, with a higher proportion of the smaller female nestlings in years of below average reproductive success. Such variation was especially pronounced in the marginal brood where a higher incidence of brood reduction allowed greater potential for sex-biased nestling mortality. In years of the highest average reproductive success, the sex ratio in the marginal brood approached equality, whereas in years of the lowest average reproductive success, more than two thirds of 8-day-old nestlings were female. Structuring the brood into core and marginal elements allowed parents to modulate both offspring number and sex under ecological uncertainty with direct consequences for population-level reproductive success. They produced fewer and less expensive fledglings in below average years and more and more expensive fledglings in above average years.     

Intra-clonal variation in the red seaweed Gracilaria chilensis

The phenotypic plasticity often found in seaweed populations has been explained only from the perspective of inter-population or inter-individual differences. However, many seaweeds grow and propagate by fragmentation of genetically identical units, each with the capacity to function on its own. If significant differences in performance exist among these supposedly identical units, such differences should be expressed upon the release and growth of these units. In this study we document two such types of variation in the red seaweed Gracilaria chilensis. Populations of sporelings, each grown under similar culture conditions and derived from carpospores shed by the same cystocarp exhibit significant differences in growth. In this species, each cystocarp develops from a simple gametic fusion, and cystocarp fusions occur too infrequently to account for the growth differences observed among recruits. In adult thalli, branches (ramets) derived from the same thallus (genet) and grown under similar conditions exhibit significant variation in growth rates and morphology. These findings have several implications. They suggest that carpospore production is not only an example of zygote amplification but that it also could increase variability among mitotically replicated units. Intra-clonal variability followed by fragmentation and re-attachment may increase intra-population variation which, in species of Gracilaria, is often larger than inter-population variation. In addition, the existence of intra-clonal variability suggests that strain selection in commercially important species may require a more continuous screening of highquality strains because of frequent genotypic or phenotypic changes in the various cultivars.     

Accounting for variation in prey selectivity by zooplankton

An ingestion-based prey selectivity function (IS) is described specifically to aid the simulation of zooplankton activity in circumstances in which selectivity varies, for example, as functions of prey palatability or abundance. In IS, the ultimate control of the rate of predation is not the external concentration of prey but the total rate of prey capture relative to predator demand. Further, “preference” is not an input (as it is for most prey selection functions) so that the selection or deselection of any or all prey can be freely altered. Hence, if required, all prey could be deselected, or the impact of inert materials or turbulence adversely affecting capture rates of any or all prey can be considered. Capture kinetics are not fixed at a common value for all items. In its basic form IS contains no more variables than other selectivity functions. The construction and use of switches to vary grazing in response to changes in abundance of alternative prey types or in prey nutritional (stoichiometric) quality is demonstrated. The advantages of using IS instead of the widely used ratio-based selectivity function is demonstrated with reference to model fits to experimental datasets for microzooplankton activity.     

Faunal variation on pelagic Sargassum

Pelagic Sargassum was collected in late summer, late winter, and early and late spring from inshore waters, the Gulf Stream and the Sargasso Sea of the Western North Atlantic Ocean. The noncolonial macrofauna was picked from the weed samples. The 34 samples contained 67 species and 11,234 individuals. The Shannon-Wiener index of diversity had a mean value of 2.419±0.177 (t.₀₅s _x ^– ) and a statistical range between 1.401 and 3.437 (t.₀₅ s). Mean diversity values were not significantly different among the various sampling series, and diversity did not vary with raft volume. High diversity values were related to an equitable distribution of species resulting from a stable environment and an area low in productivity. Species composition of the Sargassum organisms varied seasonally and geographically. Animals were more abundant in the spring than in the fall samples. Samples collected on a transect in the Gulf Stream and Sargasso Sea maintained a similar faunal composition.Contribution No. 351 from the Virginia Institute of Marine Science, Gloucester Point, Virginia, USA.From a thesis submitted to the faculty of the School of Marine Science, The College of William and Mary, in partial fulfillment of the requirements for the degree of Master of Arts in Marine Science.     

Addressing among-group variation in covariate effects using multilevel models

Multilevel models are used to model processes associated with hierarchical data structures. Despite infrequent use in the biological and environmental sciences, the use of these models with hierarchically-structured data conveys multiple advantages. These include the assessment of whether covariate effects differ among groups or clusters, and separate estimation of covariate effects by hierarchical level (thereby addressing atomistic and aggregation fallacy concerns). We illustrate these advantages using larval mayfly count data derived from annual surveys on the Mississippi River and a continuous covariate (water depth).     

Colony fusion causes within-colony variation in a parthenogenetic ant

The evolutionary stability of cooperation and altruism in colonies of social insects requires that nestmates be to some extent related. An efficient system of discrimination against non-nestmates protects the nest against unrelated conspecifics, which might exploit or parasitize the colony. The co-occurrence of unrelated individuals in mature colonies therefore is a rare event that deserves more attention. Here, we report on the relatively common incidence of colony fusion in the ant Platythyrea punctata. Workers of this ant can produce genetically identical female offspring from unfertilized eggs through thelytokous parthenogenesis. Consequently, the majority of colonies has a “clonal structure” and consists of individuals with identical multilocus genotypes. Nevertheless, field observations indicate that a surprisingly large percentage of colonies contain workers belonging to two or more different genetic lineages. Much of this genetic heterogeneity is incompatible with eventual recombination or mutation events, but instead appears to result from colony fusion or the adoption of unrelated individuals. Indeed, colonies of P. punctata from the Dominican Republic and Barbados readily merged in the laboratory and, after elimination of one of the two reproductive workers, formed stable, genetically heterogeneous colonies. We discuss the possible causes and benefits of colony fusion in natural populations.     

Bearded seal males perceive geographic variation in their trills

Geographic variation in animal vocalisations, ranging from micro- to macro-variations, has now been widely documented. These dialects can impair communication between separated groups or populations and thus may play a role in speciation processes. Although the existence of geographical variation has been already shown in some pinnipeds species, the extent to which individuals perceive it is poorly understood. Here, we studied this question in the bearded seal Erignatus barbatus, a species found across different arctic regions. As in other phocids, bearded seal males emit sophisticated acoustic displays while defending an aquatic territory during the breeding season. First, we used playbacks to demonstrate that the trill has a function in territorial defence. Second, we used synthetic trills from two distinct populations (Nunavut and Greenland), to show that bearded seal males perceived the geographic variation in the trills. Males from Greenland responded more strongly to trills from local males than those from a distant area (Nunavut). This study provides the first experimental evidence that phocid males can perceive geographic variation in their vocalisations. Acoustic analyses combined with playback experiments can help to identify distinct populations in bearded seals. This may then provide information on both strength and scale of breeding site fidelity in this and potentially other species.     

Giant pandas attend to androgen-related variation in male bleats

Although androgen-dependant traits are predicted to signal overall male quality, no study has examined the response of a nonhuman animal to variation in a known acoustic cue to male androgen levels (steroid hormones that are key drivers of male sexual behaviour). Here, we use a single-speaker approach to present male and female giant pandas with re-synthesised male bleats representing callers with high and low androgen levels. Our results revealed that male and female giant pandas had significantly greater-looking responses, spent more time pacing, and were faster to respond to playbacks of bleats simulating high androgen males. When we analysed the sexes separately, a slightly different response pattern was revealed: whereas males and females still had significantly greater-looking responses and were faster to respond to bleats simulating high androgen males, only male giant pandas tended to spend more time pacing. These findings suggest that vocal cues to male androgen levels are functionally relevant to male and female giant pandas during the breeding season, and constitute the first demonstration that a nonhuman animal could be using a vocal signal to assess male hormonal state. We go on to discuss the ecological relevance of signalling androgen levels in this species’ sexual communication and the possible application of our results to conservation breeding.