Ramet phenology and clonal architectures of the clonal sedge <Emphasis Type="Italic">Schoenoplectus americanus</Emphasis> (Pers.) Volk. ex Schinz &; R. Keller

The clonal plant Schoenoplectus americanus shows variable belowground clonal architecture as a result of producing two types of ramets: those with very long rhizomes (long rhizome ramet, LRR) and those with very short ones (short rhizome ramet, SRR). In a previous study we demonstrated that the two types of ramets are functionally specialised. The production of SRRs results in the formation of consolidated clonal patches with densely packed shoots, while the production of LRRs results in a more diffuse network of connected rhizomes with widely spaced shoots. We hypothesised that the two types of ramets would be produced at different times during the growing season because of their functional differences. The production of LRRs throughout the growing season would enable the species to continuously explore new habitats while the production of SRRs early in the growing season would enable the species to occupy and consolidate resources in available open patches. We evaluated this hypothesis through field observations in different communities with S. americanus and indeed found that SRRs were produced early in the growing season while LRRs tended to be produced over an extended period of time. Plants in high-quality environments (i.e. higher light conditions) produced more SRRs, and these were formed early in the growing season. In contrast, plants in low-quality environments produced more LRRs, and these were formed continuously over the growing season. We also observed that the shoot longevity was greater for SRR. In high-quality patches, the production of the lower cost SRRs results in a more rapid occupancy of open spaces; in lower quality patches, the production of LRRs throughout the growing season enables plants to explore the immediate environment for higher quality patches.     

Characterization of microsatellite loci in Schoenoplectus americanus (Cyperaceae)

Schoenoplectus americanus is a model organism for studying ecological and ecosystem responses of salt marsh plant communities to global climate change. Here we characterize 16 microsatellite loci in S. americanus to facilitate studies on the genetic basis of phenotypic responses to changing climate conditions such as elevated atmospheric carbon dioxide. Most loci also amplified in the morphologically similar sister species, Schoenoplectus pungens. Five loci exhibited species‐specific alleles or distinct allelic size distributions that discriminate S. americanus from S. pungens.     

Phytoremediation of two types of sediment contaminated with Zn by Schoenoplectus americanus

The effect of different sediments on growth, Zn uptake, Zn plant distribution, and morphometric variables of Schoenoplectus americanus were investigated under controlled conditions. Two types of sediments were assayed: from a large natural levee (LS) and alluvial sediments (AS), the former with lower organic matter (OM) and nutrients content than AS, without and with added Zn (2500 microg Zn/g air-dry sediment). Zinc partition in sediment was determined. Increases in water conductivity and Zn concentrations in water and sediments were observed in artificially contaminated treatments. Plants showed a lower above ground growth rate, height, and width of shoots, and a higher Zn concentration in shoots and rhizomes. In the contaminated treatments, AS treatment showed lower Zn concentration in water and higher Zn concentration in sediments (total, exchangeable, and OM fractions) than LS treatment, due to Zn displacement from floodwater to sediments. The presence of a high level of OM and nutrients also increased aboveground biomass growth, whereas it decreased Zn concentration in shoots. Although the translocation factor increased with Zn addition, it was lower in AS treatment Sediments of AS treatments are a suitable environment for growth of S. americanus, which partially compensates the toxic effects of Zn. Our results provide an encouraging basis for planning larger scale experiments to test the role of OM and nutrients in improving phytoremediation.     

Integration in the clonal plant <Emphasis Type="Italic">Eriophorum angustifolium</Emphasis>: an experiment with a three-member-clonal system in a patchy environment

A clonal plant in heterogeneous environments is usually expected to profit from resource exchange via a clonal network where ramets placed in contrasting environments can specialise so to acquire the most abundant resources. An experiment was designed using the three member clonal system of Eriophorum angustifolium, which consisted of one parent ramet growing in a resource poor environment and two offspring: one was limited in growth by nutrients while the other was light limited; the contrast in availability of limited resources between the offspring ramets was high, medium or none, with the system either connected or severed. The total resource availability was the same in all treatments. We proposed four possible scenarios for the system: offspring ramets will share resources via the deficient parent ramet, and the whole clone will profit from the contrasting environment (scenario 1); offspring ramets will support exclusively the parent ramet, and the whole clone will profit from a homogeneous environment (scenario 2); offspring ramets will stop the export of the limiting resource to the parent ramet, with split and connected treatments not differing (scenario 3); and offspring ramets will exhaust the carbon stored in the biomass of the parental ramet; offspring ramet will profit from connection (scenario 4). In the experiment, the limiting resources were sent to the strongest sink (scenario 2). The parent ramet growing in a deficient environment received the highest support in the treatment where both offspring ramets were growing in the same conditions (no-contrast treatment). Production of new shoots, but not biomass of whole clone, was supported in a homogenous environment. The experiment revealed that multiple stresses might prohibit free exchange of limiting resources via the clonal network and supports the idea that experimental studies on more complex clones are essential for understanding the costs and benefits of clonal growth.     

Ropalidia rufoplagiata: a polistine wasp society probably lacking permanent reproductive division of labour

Summary Ropalidia rufoplagiata Cameron (Hymenoptera: Vespidae), a polistine species from penisular India, appears to be unique among all known primitively eusocial wasps. A total of 33 out of 46 identified females from an observed colony were found to oviposit on 1–17 occasions. No single predominant egg-layer could be identified during the 45-day period. Of the 17 dissected egg-layers, 12 were mated. All egg-layers showed several oviposition-related behavioural patterns including systematic, but indiscriminate, cannibalism of eggs and larvae, cleaning of empty cells, and guarding of freshly-laid eggs. There was no correlation between the egg-laying activity of the females (whether mated or not), oophagy, and their position in the dominance hierarchy. All nest-maintenance activities were performed exclusively by the egg-layers, while the non-egg-layers were mainly involved in the extranidal task of foraging. No significant morphometric differences between egg-layers and foragers could be discerned. Almost all the older individuals in the colony were egg-layers, while foragers were mainly younger animals. Such a temporal differentiation in reproductive labour suggests the absence of a permanent reproductive caste in this species.     

Estimation of larval dry weight of Chaoborus americanus

A method for estimating larval dry weight as a function of tracheal front air sac length and instar was developed for second, third, and fourth instar Chaoborus americanus using linear regression analysis. The following equation: where W = larval dry weight (μg), L = front air sac length (mm), a = the y-intercept, and b = the slope provided accurate estimates of larval dry weight for three C. americanus populations. No detectable change in size occurred in the air sacs of larvae preserved for one year in 5% formalin. This indicates that air sacs can be used to accurately estimate larval dry weight for preserved as well as living specimens.     

Progress towards characterization of a predator/prey kairomone: Daphnia pulex and Chaoborus americanus

Predatory larvae of the midge Chaoborus americanus induce an antipredator defense (neckteeth) in prey individuals of the cladoceran Daphnia pulex. The signal for presence of predator is a water-soluble chemical. We provide evidence that this kairomone originates in the intestinal tract of the predator. The active compound is an organic molecule of intermediate polarity which is heat stable and partially destroyed by acid and base digestion. It is stable to digestion by the general peptidase Pronase. Hydroxyl groups, but not primary amines, carbonyls or thiols are essential to activity. Low-pressure liquid chromatography on a reverse-phase silica gel (Amicon Matrex C-18) column suggests there may be more than one active component.     

Effects of simulated feeding by snow geese on Scirpus americanus rhizomes

Summary We simulated the feeding of Greater Snow Geese (Chen caerulescens atlantica) on the rhizomes of three-square bulrush (Scirpus americanus) in a tidal marsh along the St. Lawrence River estuary in Québec. During the spring staging period, aboveground biomass is unavailable and geese feed solely on rhizomes and overwintering buds. An experiment was designed to test the effect of three factors on subsequent growth of Scirpus: the intensity of removal (3 to 77% removal of belowground biomass), the number of bites (1, 2 or 3 sections removed) and the number of adventitious buds removed (1, 2 or 3). Rhizomes were dug out in May, treated and transplanted into 85-1 basins sunk in the marsh and filled with marsh soil freed of all plant material. Growth was observed weekly until the end of the growing season in August. Shoots and rhizomes were then collected, dried and weighed to obtain biomass estimates. The net above- and belowground production of Scirpus was inversely related to the initial rhizome biomass removed. At a high level of removal (>35%), the cumulative number of shoots was significantly reduced as early as two weeks after transplantation. The relative reduction in production of the treated rhizomes compared to the control plants was also related to the intensity of removal. An increased number of bites reduced production and the removal of an increased number of adventitious buds further amplified the effect of removal on rhizome production. These experimental results show that even low intensity of feeding by Snow Geese can reduce the production of Scirpus marshes.     

Geometry explains the benefits of division of labour in a leafcutter ant

Many ant species have morphologically distinct worker sub-castes. This presumably increases colony efficiency and is thought to be optimized by natural selection. Optimality arguments are, however, often lacking in detail. In ants, the benefits of having workers in a range of sizes have rarely been explained mechanistically. In Atta leafcutter ants, large workers specialize in defence and also cut fruit. Fruit is soft and can be cut by smaller workers. Why, therefore, are large workers involved? According to the geometry hypothesis, cutting large pieces from three-dimensional objects like fruit is enhanced by longer mandibles. By contrast, long mandibles are not needed to cut leaves that are effectively two-dimensional. Our results from Atta laevigata support three predictions from the geometry hypothesis. First, larger workers cut larger fruit pieces. Second, the effect of large size is greater in cutting fruit than leaves. Third, the size of fruit pieces cut increases approximately in proportion to the cube of mandible length. Our results are a novel mechanistic example of how size variation among worker ants enhances division of labour.     

Mitotic and meiotic chromosomes of the American lobster Homarus americanus (Nephropidae, Decapoda)

The chromosomes of H. americanus have been characterised by C-banding, fluorochrome banding and restriction endonuclease banding. Thanks to these techniques, it has been possible to identify mitotic and meiotic figures clearly and to study the distribution and structure of heterochromatic regions. Moreover, we have identified small supernumerary chromosomes, variable in number and often asynaptic in first meiotic metaphase.     

Temporal and spatial patterns of wax secretion and related behavior in the division of labour of the honeybee (Apis mellifera capensis)

Summary Current ethograms for honeybee behavior are hypothesized to consist of age-related cohorts of workers having a high probability of performing a small set of related tasks in a restricted portion of the nest. Wax secretion and wax working related activities were assayed in this light. It is shown that wax secretion is significantly affected by worker age and that bees between 3 and 21 days old form such a cohort. Comb building festoons, previously thought to be the site of wax secretion, represent only a small fraction of newly secreted wax in the nest. Wax secretion remains constant relative to age in the cohort but varies significantly with season as does bee participation in festooning behavior. Wax secretion and wax working are both definable in terms of time and space in the nest, the relative probability of activity changing with season. Secretion itself is constrained by the cyclical activity of the underlying wax gland complex.     

Characterization of 29 tetranucleotide microsatellite loci in black bear (Ursus americanus) for use in forensic and population applications

We developed 29 forensic quality tetranucleotide microsatellite loci for the identification of individual black bear (Ursus americanus). Allele number averaged 5.35 and ranged from 3 to 11. The eight markers selected for forensic use permit the identification and exclusion of individual bear with a probability of identity of approximately 8.1 × 10⁻⁸ (approximately one per 12 million).     

Two types of division of labour in clonal plants: benefits, costs and constraints

Clonal plants spread vegetatively within their habitats by forming rooted ramets on stolons or rhizomes. Each of these ramets is capable of an independent existence after establishment. Nevertheless, ramets remain physically connected by stolon or rhizome internodes for variable periods of time, thereby allowing for resource movement and signal transduction within clones.Interconnected ramets of clonal plants, though potentially independent and totipotent, can specialize functionally in the performance of limited numbers of tasks such as the uptake of resources from above- vs below-ground sources, carbohydrate storage, vegetative spread and sexual reproduction. Such specialization and cooperation is comparable to a division of labour in economic systems or in colonies of social animals. The ecological significance of division of labour in clonal plants may be found in the increased efficiency of entire clones in exploiting their environments.Two different types of division of labour in clonal plants will be discussed in this review. The first type is an environmentally-induced specialization of ramets in the uptake of locally abundant resources (plastic division of labour), which can be found in several stoloniferous species. Evidence exists that this response increases resource uptake in spatially heterogeneous environments. The second type of division of labour, which occurs mainly in rhizomatous species, relates to a developmentally-programmed specialization and cooperation between interconnected ramets. This response pattern is thought to enhance plant performance by restricting the number of tasks for individual ramets and thereby significantly increasing the efficiency of task performance. In some plants, such an inherent division of labour is likely to contribute to nutrient extraction from poor and unpredictably variable sources.In this article not only benefits but also potential costs and constraints on division of labour in clonal plants are shown. The aim is to provide a review of existing knowledge and to develop concepts and hypotheses for future research.     

A Review of Hypotheses of Decline of the Endangered American Burying Beetle (Silphidae: Nicrophorus americanus Olivier)

The largest species of North American Nicrophorus (Coleoptera: Silphidae), N. americanus, was placed on the US federal list of endangered species in 1989. This paper reviews literature bearing on eight hypotheses that attempt to explain the dramatic decline of this species over 90% of its former range. What is known regarding each hypothesis is separated from what remains to be investigated. We find that although progress has been made during the past 12 years, even the most well supported hypothesis requires a number of important studies to be completed or extended before we can confidently explain the decline of this species and predict the success of conservation efforts.     

The evolutionary history of division of labour

Functional specialization, or division of labour (DOL), of parts within organisms and colonies is common in most multi-cellular, colonial and social organisms, but it is far from ubiquitous. Several mechanisms have been proposed to explain the evolutionary origins of DOL; the basic feature common to all of them is that functional differences can arise easily. These mechanisms cannot explain the many groups of colonial and social animals that exhibit no DOL despite up to 500 million years of evolution. Here, I propose a new hypothesis, based on a multi-level selection theory, which predicts that a reproductive DOL is required to evolve prior to subsequent functional specialization. I test this hypothesis using a dataset consisting of the type of DOL for living and extinct colonial and social animals. The frequency distribution of DOL and the sequence of its acquisition confirm that reproductive specialization evolves prior to functional specialization. A corollary of this hypothesis is observed in colonial, social and also within multi-cellular organisms; those species without a reproductive DOL have a smaller range of internal variation, in terms of the number of polymorphs or cell types, than species with a reproductive DOL.     

Life expectancy and onset of foraging in the honeybee (Apis mellifera)

Predictions that precocious foraging in honeybee workers is a result of shortening of their life expectancy were tested in both laboratory and field experiments. In the laboratory experiment, we assessed the impact of anaesthesia with CO₂ and infection with Nosema apis on the lifespan of workers. In the field experiment, the age at onset of foraging was observed in groups of workers with different expected lifespans. In both the experiments, workers originating from one queen inseminated with the semen of one drone were divided into five groups. The first group was anaesthetized with CO₂ on the first day of life. Workers from the other three groups were individually inoculated with a constant number of N. apis spores on the 1st, 6th and 11th days of life. Workers from the fifth control group were neither treated with CO₂ nor inoculated in any way. Both the laboratory and field experiments revealed that anaesthetized and infected workers had shorter expected lifespans compared to control bees. Amongst infected workers, those inoculated earlier in life survived for a significantly shorter period of time in comparison to those infected later in life. In agreement with the expectation, the field experiment showed that anaesthetized and infected workers with shorter expected lifespan start foraging earlier than control workers. Amongst the infected workers, age at inoculation correlated with age at onset of foraging. This means that short-lived workers complete safe nest tasks and begin riskier foraging earlier in life. Our results provide a strong support for the hypothesis that the division of labour in eusocial insects is a consequence of the different expected worker lifespan and different risk associated with their tasks; however, they do not contradict other existing explanations.