Some statistical and conceptual issues in the detection of low-level environmental pollutants

With increasing concern over chemicals that are potential health hazards at low levels, determination of limits of detection have undergone considerable scrutiny. Most traditional detection limit estimators suffer from extensive statistical and/or conceptual limitations. In this paper, traditional detection limit estimators are described and critically evaluated. Using the terminology of Currie (1968), methods are categorized into decision limits versus detection limits. The methods are further categorized into single concentration design versus calibration design methodologies. While the single concentration design methods are useful for fixing ideas and clarifying definitions, they are shown to be extremely limited in practice since dependence of variability on concentration can neither be estimated or incorporated. Calibration-based detection limit estimators are described, compared and contrasted. Generalizations to non-constant variance, multiple future detection decisions and simultaneous control of Type I and II errors are provided. The various calibration-based methods are illustrated using real data and experimental design issues for detection limit studies discussed.     

Subordinate males are more likely to help if unrelated to the breeding female in cooperatively breeding white-browed scrubwrens

Subordinates in communally breeding groups of birds usually help to provision nestlings, but in some species helping-at-the-nest is facultative. In species in which groups usually contain relatives, subordinates either always feed young or are more likely to do so when breeding dominants are close relatives, suggesting that benefits of helping collateral kin are important. In other species, adult group members are unrelated to each other and males may only feed young if they have gained paternity, showing that cooperation is related to the mating system. The white-browed scrubwren, Sericornis frontalis, is a communally breeding species in which most groups consist of a simple pair or a dominant pair with a subordinate male. Subordinate males either fed nestlings in a given nest at a rate comparable to the dominants, or did not feed them at all. Breeding groups usually formed through natal philopatry of males, so that about 80% of subordinates were closely related to one or both members of the dominant pair. However, because of death and dispersal, 54% of subordinates were unrelated to the resident female. Although subordinates with their mother fed nestlings in 48% of cases, they fed offspring in 75% of cases if their mother had been replaced by an unrelated female, suggesting that their decision to help is influenced by the opportunity to mate with the female. Supporting this conclusion, relatedness to the dominant male did not affect subordinate behaviour, and genetic studies showed that subordinates often gained paternity if unrelated to the female. Thus, paradoxically, provisioning nestlings is related to the opportunity for mating in a society in which there is natal philopatry and subordinates are usually related to one or both members of the dominant pair. Received: 25 January 1997 / Accepted after revision: 1 May 1997     

Decentralized regulation of a common property renewable resource industry with irreversible investment

The effectiveness of output controls for rationalizing a common property renewable resource has been called into question by the theoretical work of J. R. Gould [Economica, Nov., 383–402 (1972)]. A proper examination of this question requires an intertemporal analysis, one that takes into account that asymmetries between persistent factors of production (“immaleable capital”) and factors that are instantaneously consumed (“labor” or ldharvest effort”). We present here a nonlinear intertemporal model of a renewable resource industry, under conditions of irreversible capital investment, and undertake to analyze its dynamics, both at open access and under centralized optimal management. We then examine the theoretical possibility of decentralized regulation by Pigouvian taxes, and reconsider the proposition of Gould.     

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     

Potential effects of forest policies on terrestrial biodiversity in a multi-ownership province.

We used spatial simulation models to evaluate how current and two alternative policies might affect potential biodiversity over 100 years in the Coast Ranges Physiographic Province of Oregon. This 2.3-million-ha province is characterized by a diversity of public and private forest owners, and a wide range of forest policy and management objectives. We evaluated habitat availability for seven focal species representing different life histories. We also examined how policies affected old-growth stand structure, age distributions relative to the historical range of variability, and landscape patterns of forest types. Under the current policy scenario, the area of habitat for old-growth forest structure and associated species increased over time, the habitat for some early-successional associates remained stable, and the area of hardwood vegetation and diverse early-successional stages declined. The province is projected to move toward but not reach the historical range of variation of forest age classes that may have occurred under the wildfire regimes of the pre-Euroamerican settlement period. Ownership explained much of the pattern of biodiversity in the province, and under the current policy scenario, its effect increased over time as the landscape diverged into highly contrasting forest structures and ages. Patch type diversity declined slightly overall but declined strongly within ownerships. Most of the modeled change in biodiversity over time resulted from policies on public forest lands that were intended to increase the area of late-successional forests and species. One of the alternative policies, increased retention of wildlife trees on private lands, reduced the contrast between ownerships and increased habitat availability over time for both early- and late-successional species. Analysis of another alternative, stopping thinning of plantations on federal lands, indicated that current thinning regimes improve habitat for the Olive-sided Flycatcher, but the no-thinning alternative had no effect on the habitat scores for the late-successional species in the 100-year simulation. A comparison of indicators of biological diversity suggests that using focal species and forest structural measures can provide complementary information on biodiversity. The multi-ownership perspective provided a more complete synthesis of province-wide biodiversity patterns than assessments based on single ownerships.     

Deception by helpers in cooperatively breeding white-winged choughs and its experimental manipulation

White-winged choughs (Corcorax melanorhamphos) are obligate cooperative breeders, living in groups which may contain up to 20 birds. Although breeding is dominated by a single pair, all birds contribute to rearing young, including the provisioning of nestlings. However, some birds which have carried food to the nest, even to the point of placing the food in the gaping mouth of a nestling, consume the food themselves rather than provision the nestlings. Birds which fail to feed nestlings are typically young, and are only likely to fail to deliver food when they cannot be observed by other group members. Birds which have just failed to deliver food are more likely to engage in alternative helping behaviours such as allopreening the nestlings than are helpers which have just delivered food in the conventional manner. Failure to deliver food is almost eliminated when foraging constraints are experimentally reduced by supplemental feeding of the group. Collectively these observations suggest that young white-winged choughs act deceptively by simulating helping behaviours without sacrificing food supplies. Received: 24 January 1997 / Accepted after revision: 6 June 1997     

Influence of kinship on helping behavior in Galápagos mockingbirds

Summary The social organization of the Galápagos mockingbird (Nesomimus parvulus) in unusual in that groups frequently include more than one breeding pair (plural breeding), and helping behavior is flexible: some birds neither breed nor help, while others do both. To investigate the influence of kinship on helping behavior, I categorized each bird as a helper or non-helper with respect to each nest within its group where it had an opportunity to help. The incidence of helping varied with relatedness: more birds helped when nestlings available to be fed were close relatives than when not. This result was independent of a higher incidence of helping among males than among females and of variation with age among males. Proportionally more nonbreeding than breeding males helped, but breeding and nonbreeding females helped equally infrequently; breeders helped most often after their own nests failed. The incidence of helping was highest among birds with opportunities to feed offspring of breeders that had fed the potential helper as a nestling, suggesting a mechanism for kin discrimination based on associative learning. Juveniles with opportunities to choose among alternative recipients preferentially fed closely related nestlings, but insufficient information was available to determine if adults also did so. Kinship did not influence the rate at which nestlings were fed by helpers. Juveniles fed nestlings at lower rates than did adult helpers, but helping effort was otherwise unaffected by age, sex, or relatedness. Limitation of help to former feeders functions as a mechanism for directing aid to relatives in a plural breeding system where degrees of kinship vary among potential recipients within the same group.     

Local adaptation of immunity against a trematode parasite in marine amphipod populations

Resources allocated to defence against parasites are not available for investment in other functions such as growth or reproduction, resulting in trade-offs between different components of an organism’s fitness. In balancing the cost of infection and the cost of immunity, selection should only favour individuals that allocate more energy to resistance and immune responses in populations regularly exposed to debilitating parasites. Here, we compare the ability of amphipods, Paracalliope novizealandiae, to (1) avoid becoming infected and (2) to respond to infection by encapsulating and melanizing parasites, between two natural populations exposed to different risk of parasitism. One population faces high levels of infection by the debilitating trematode parasite Maritrema novaezealandensis, whereas the other population is not parasitised by this trematode nor by any other parasite. Under controlled experimental conditions, with exposure to a standardized dose of parasites, amphipods from the parasite-free population acquired significantly more parasites than those from the population regularly experiencing infection. Furthermore, a lower frequency of amphipods from the parasite-free population succeeded at melanizing (and thus killing) parasites, and they melanized a lower percentage of parasites on average, than amphipods from the parasitised population. These differences persist when individual factors, such as amphipod sex or body length, are taken into account as potential confounding variables. These results support the existence of local adaptation against parasites: an amphipod population that never experiences trematode infections is less capable of resisting infection, both in terms of its first line of defence (avoiding infection) and a later line of defence (fighting parasites following infection), than a population regularly exposed to infection.     

Isotopic evidence for in-lake production of accumulating nitrate in Lake Superior.

A century-long increase in nitrate (NO3-) in the water column of Lake Superior is a classic example of recent nitrogen accumulation in ecosystems, but its cause and relationship to historical NO3- deposition is unknown. We used stable isotope ratios of oxygen and nitrogen in nitrate (delta18O-NO3 and delta15N-NO3) to examine its sources in this large lake, which represents 10% of the world's surficial liquid freshwater. The most parsimonious hypothesis to explain the rise in NO3- is that the lake is accruing NO3- deposited directly on the lake surface because it is too unproductive to completely assimilate all of it. Data for delta18O-NO3 in external sources and the water column, however, are inconsistent with this hypothesis. Instead, the isotopic evidence indicates strongly that the accumulating NO3- is almost entirely derived from nitrification occurring within the lake. While increases in atmospheric deposition of NO3- may have played a role in its buildup in the lake, other factors such as increases in NH4+ and dissolved organic nitrogen inputs from precipitation or rivers, increases in nitrogen fluxes from the sediments, and decreases in burial rates must also be considered as potential drivers of rising NO3-. The sustained accumulation of nitrogen in Lake Superior is thus more complex and incompletely understood than previously assumed.     

Ovariole number—a predictor of differential reproductive success among worker subfamilies in queenless honeybee (Apis mellifera L.) colonies

A honeybee queen normally mates with 10–20 drones, and reproductive conflicts may arise among a colony’s different worker patrilines, especially after a colony has lost its single queen and the workers commence egg laying. In this study, we employed microsatellite markers to study aspects of worker reproductive competition in two queenless Africanized honeybee colonies. First, we determined whether there was a bias among worker patrilines in their maternity of drones and, second, we asked whether this bias could be attributed to differences in the degree of ovary activation of workers. Third, we relate these behavioral and physiological factors to ontogenetic differences between workers with respect to ovariole number. Workers from each of three (colony A) and one (colony B) patrilineal genotypes represented less than 6% of the worker population, yet each produced at least 13% of the drones in a colony, and collectively they produced 73% of the drones. Workers representing these genotypes also had more developed follicles and a greater number of ovarioles per ovary. Across all workers, ovariole development and number were closely correlated. This suggests a strong effect of worker genotype on the development of the ovary already in the postembryonic stages and sets a precedent to adult fertility, so that “workers are not born equal”. We hypothesize a frequency-dependent or “rare patriline” advantage to queenless workers over the parentage of males and discuss the maintenance of genetic variance in the reproductive capacity of workers.Electronic supplementary material Supplementary material is available for this article at and is accessible for authorized users.