Sexual dimorphism and cost of reproduction in the dioecious shrub Lindera benzoin (Lauraceae)>

We examined sexual dimorphism in reproductive allocation and its effects on growth and subsequent reproduction in a natural population of the dioecious woody shrub Lindera benzoin (L.) Blume. In addition to comparing natural patterns of growth and reproduction in a marked population of 251 females and 87 males, we experimentally examined the effect of reduced reproduction on future growth and reproduction in female plants and examined sexual dimorphism in carbon/nutrient balance. Our results suggest that females of L. benzoin bear greater reproductive costs in terms of both biomass and minerals. These costs were measurable in terms of current biomass and allocation of nitrogen to reproduction, as well as subsequent growth, reproduction, and tissue carbon/nutrient balance. Based upon the results of a fruit-thinning experiment and path analysis, fruit production in 1991 appeared to have direct negative effects on fruit production in 1992, an effect that was not necessarily mediated through effects on plant growth. We discuss our results in the context of other intrinsic and extrinsic factors that can influence growth, reproductive costs, and mortality in this species.     

Reproductive allocation in hermaphrodite and female plants of Sidalcea oregana SSP. spicata (Malvaceae) using four currencies

Reproductive allocation was investigated in female and hermaphrodite plants of gynodioecious Sidalcea oregana ssp. spicata. Total reproductive investment and partitioning of that investment was documented at the level of whole plants in terms of four ecologically relevant currencies: biomass, nitrogen, phosphorus, and potassium. Nutrient augmentations in the field confirmed that nutrients were limiting plant vegetative growth and propensity to flower; thus the use of these nutrients as currency was appropriate. Once the effects of plant size were removed, the sex morphs allocated similar total amounts of biomass, nitrogen, phosphorus, and potassium to reproduction, but partitioned those differentially. For any given individual size, females allocated larger proportions of their reproductive resource budgets to seeds. Hermaphrodites' reproductive investment in pollen and flowers was allocated at the expense of allocation to seeds. These data are relevant to the evolution of gynodioecy from hermaphroditism and support the hypothesis that females reallocate resources not spent on pollen to seeds.     

Functional dietary response of Asian black bears to changes in sika deer density

Omnivores are generally opportunistic foragers and have a flexible dietary response to resource abundance and availability. Their populations may consist of individuals that differ from each other in terms of their trophic positions, which implies that the dietary response to resource fluctuations differs within a population. We investigated how changes in the abundance of sika deer (Cervus nippon) affected dietary variation and body condition in the Asian black bear (Ursus thibetanus). We used fecal analysis, nitrogen stable isotopes (δ¹⁵N), and body measurements to determine whether the variation in dietary meat content of Asian black bears is positively related to variations in the density of the sika deer population, whether male bears have a higher trophic position compared to females, and whether dietary meat content is positively related with body mass or body condition of bears. We found a positive correlation between the occurrence of deer remains in bear feces and deer density, suggesting that bears change their diet in response to temporal changes in deer density. Male bears had higher δ¹⁵N values than females, and neither values varied when deer density decreased. Males selectively consumed deer after a reduction in deer density, whereas females consistently consumed more plant-based diet. The δ¹⁵N values were positively related with body mass of adult (>4 yr) bears but had no relationship with body condition of bears of either sex or any age class. Deer seem to be an important food source for large adult males, which have an advantage in mating. Thus, increasing herbivore abundance and availability altered the foraging strategy of Asian black bears, but the importance of herbivore on bear diet differs within a population.     

Sexual differences in gas exchange and response to environmental stress in dioecious Silene latifolia (Caryophyllaceae)

Females of dioecious species usually have higher reproductive effort than males because they produce fruits in addition to flowers. Since females have higher reproductive effort, they are expected to be more negatively affected than males by low resource availability. We tested that assumption by growing females and males of Silene latifolia under low levels of light, water, nitrogen, phosphorus, and potassium. Gas exchange of the sexes did not respond differently to low resource availability; higher female reproductive effort relative to males did not differentially affect their ability to assimilate carbon. However, male photosynthesis rates and stomatal conductances were slightly, but consistently, higher than those of females. The intersexual difference in photosynthesis rate may be a proximate result of reproduction if females translocate nutrients, particularly nitrogen, from their leaves to developing fruits. Alternatively (or perhaps additionally), higher male photosynthesis and stomatal conductances relative to females may be the ultimate result of sexual selection. This could be the case if 1) reproductive effort as estimated by biomass allocation is misleading and males actually invest more in reproduction than females, or 2) females experience stronger selection than males to conserve water late in the growing season, when soil moisture is likely to be low but females need to complete fruit maturation. Our results indicated that females had slightly lower leaf nitrogen but higher photosynthetic water-use efficiency than males, so it is possible that both proximate and ultimate factors are operating simultaneously to cause lower female photosynthesis rates.     

Nitrogen uptake and preference in a forest understory following invasion by an exotic grass

Plant–soil interactions have been proposed as a causative mechanism explaining how invasive plant species impact ecosystem processes. We evaluate whether an invasive plant influences plant and soil-microbe acquisition of nitrogen to elucidate the mechanistic pathways by which invaders might alter N availability. Using a ¹⁵N tracer, we quantify differences in nitrogen uptake and allocation in communities with and without Microstegium vimineum, a shade-tolerant, C₄ grass that is rapidly invading the understories of eastern US deciduous forests. We further investigate if plants or the microbial biomass exhibit preferences for certain nitrogen forms (glycine, nitrate, and ammonium) to gain insight into nitrogen partitioning in invaded communities. Understory native plants and M. vimineum took up similar amounts of added nitrogen but allocated it differently, with native plants allocating primarily to roots and M. vimineum allocating most nitrogen to shoots. Plant nitrogen uptake was higher in invaded communities due primarily to the increase in understory biomass when M. vimineum was present, but for the microbial biomass, nitrogen uptake did not vary with invasion status. This translated to a significant reduction (P < 0.001) in the ratio of microbial biomass to plant biomass nitrogen uptake, which suggests that, although the demand for nitrogen has intensified, microbes continue to be effective nitrogen competitors. The microbial biomass exhibited a strong preference for ammonium over glycine and nitrate, regardless of invasion status. By comparison, native plants showed no nitrogen preferences and M. vimineum preferred inorganic nitrogen species. We interpret our findings as evidence that invasion by M. vimineum leads to changes in the partitioning of nitrogen above and belowground in forest understories, and to decreases in the microbial biomass, but it does not affect the outcome of plant–microbe–nitrogen interactions, possibly due to functional shifts in the microbial community as a result of invasion.     

Sexual differences in biomass and nutrient allocation of first-year Silene dioica plants

Reproductive and somatic biomass, nitrogen (N), and phosphorus (P) pools were compared between females and males in 1st-year plants of Silene dioica. We estimated irretrievable resources allocated to seeds, pollen, flowers, and unrecovered summer leaf investment by collecting plant parts at abscission throughout the season. At the end of the season, we determined resources lost through senescent stems and autumn leaf turnover and resources stored in perennial roots and overwintering buds. Sexual differences in allocation patterns depended on the resource used for comparison, and whether absolute or proportional resource pools were assessed. Total resource pools in terms of biomass and N were similar for females and males. However, male plants acquired relatively more P. The proportional reproductive investment, i.e., reproductive effort, was similar for males and females in terms of biomass and N. In terms of P, male reproductive effort was higher. There was no difference between sexes in the proportional and relative biomass allocated to perennial roots and overwintering buds. However, in terms of absolute and relative N allocation to below-ground parts, females had larger reserves than males. Females, moreover, had a larger proportion of their P in below-ground parts. However, as male total P pools were larger, absolute P reserves did not differ between sexes. The high reproductive effort and N depletion of below-ground parts in males resulted largely from higher flower production compared to females. In females, seeds were the major component of reproductive effort. These results show that if biomass and nutrient allocation are assessed in parallel for dioecious plants, we obtain a more complete view of their sexual differences. Received: 07 May 1998 / Accepted: 30 October 1998     

Roots, shoots and reproduction: sexual dimorphism in size and costs of reproductive allocation in an annual herb

Females tend to be smaller than males in woody dioecious plant species, but they tend to be larger in herbs. The smaller size of females in woody species has been attributed to higher reproductive costs, yet no satisfactory explanation has been provided for their larger size in herbs. Because herbs have higher nitrogen concentrations in their tissues than woody plants, and because pollen is particularly rich in nitrogen, we predicted that male growth would be more compromised by reproduction than female growth. To test this hypothesis, we conducted three experiments on the annual dioecious herb Mercurialis annua. First, we compared the timing of reproduction between males and females and found that males started flowering earlier than females; early flowering is expected to compromise growth more than later flowering. Second, we compared plants allowed to flower with those prevented from flowering by experimental debudding and found that males incurred a higher reproductive cost than females in terms of both biomass and, particularly, nitrogen. Third, we grew plants under varying levels of nitrogen availability and found that although sexual size dimorphism was unaffected by nitrogen, females, but not males, decreased their relative allocation to both roots and reproduction under high nitrogen availability. We propose that males deal with the high cost of pollen production in terms of nitrogen by allocating biomass to nitrogen-harvesting roots, whereas females pay for carbon-rich seeds and fruits by investing in photosynthetic organs. Sexual dimorphism would thus seem to be the outcome of allocation to above- versus below-ground sinks that supply resources (carbon versus nitrogen) limiting the female and male reproduction differentially.     

Nutrient limitation and plant species composition in temperate salt marshes

Addition of inorganic nitrogen, phosphorus and potassium in a factorial design in two ungrazed Wadden-Sea salt marshes at low and high elevations showed that nitrogen was the limiting nutrient. No effects of nutrient addition were detected in the 1st year, probably due to a considerable rainfall deficit during the growing season. In the 2nd year, which was more humid, only nitrogen addition caused significant effects in both the low salt marsh dominated by Puccinellia maritima and the high marsh dominated by Festuca rubra. No two-way or three-way interactions with phosphorus or potassium were found. In the low marsh, nitrogen addition had a negative effect on the biomass of Puccinellia, but a positive effect on the biomass of Suaeda maritima and on the total above-ground biomass. Puccinellia was replaced by Suaeda after nitrogen addition, due to shading. In the high salt marsh, no significant effects of fertilizer application on total above-ground biomass were found, due to the weak response of the dominant species Festuca rubra, which accounted for 95% of total biomass. The biomass of Spergularia maritima increased, however, as a response to nitrogen addition.The shoot length of Festuca was positively affected by nitrogen fertilization. It is suggested that stands of Festuca reached maximal biomass at the study site without fertilization and that its growth was probably limited by self-shading. Received: 22 September 1996 / Accepted: 5 April 1997     

DIFFERENCES IN BIOMASS ALLOCATION TO REPRODUCTIVE AND VEGETATIVE STRUCTURES OF MALE AND FEMALE PLANTS OF A DIOECIOUS,PERENNIAL HERB,SILENE ALBA (MILLER) KRAUSE

The pattern of biomass allocation of males and females and the sex ratio and growth characteristics of plants from three seed-size classes in Silene alba were investigated in a greenhouse study. Seed size significantly affected adult plant size and flower production of both male and female plants, but there was no significant difference in the proportion of males and females emerging in three seed-size categories. Male and female plants differed in the proportion of total biomass allocated to vegetative and reproductive structures and these differences were consistent across all seed-size categories. Males allocated a greater proportion of their biomass to flowers than did females. Female reproductive effort was dependent upon the percentage of flowers producing mature capsules. Only females with greater than 20% fruit set have a higher reproductive expenditure than males. Consequently, female expenditure is potentially greater than males, but is spread out over a longer portion of the growing season. This difference in the timing of reproductive expenditures by males and females allows females to allocate more biomass to growth during the early flowering period and may therefore account for the common pattern in herbaceous perennial dioecious species in which adult females are larger than adult males.     

Effects of European Earthworm Invasion on Soil Characteristics in Northern Hardwood Forests of Minnesota, USA

European earthworms are colonizing worm-free hardwood forests across North America. Leading edges of earthworm invasion in forests of northern Minnesota provide a rare opportunity to document changes in soil characteristics as earthworm invasions are occurring. Across leading edges of earthworm invasion in four northern hardwood stands, increasing total earthworm biomass was associated with rapid disappearance of the O horizon. Concurrently, the thickness, bulk density and total soil organic matter content of the A horizon increased, and it’s percent organic matter and fine root density decreased. Different earthworm species assemblages influenced the magnitude and type of change in these soil parameters. Soil N and P availability were lower in plots with high earthworm biomass compared to plots with low worm biomass. Decreases in soil nitrogen availability associated with high earthworm biomass were reflected in decreased foliar nitrogen content for Carex pensylvanica, Acer saccharum and Asarum canadense but increased foliar N for Athyrium felix-femina. Overall, high earthworm biomass resulted in increased foliar carbon to nitrogen ratios. The effects of earthworm species assemblages on forest soil properties are related to their feeding and burrowing habits in addition to effects related to total biomass. The potential for large ecosystem consequences following exotic earthworm invasion has only recently been recognized by forest ecologists. In the face of rapid change and multiple pressures on native forest ecosystems, the impacts of earthworm invasion on forest soil structure and function must be considered.     

植株密度和氮素互作对垂穗披碱草生物量分配的影响

垂穗披碱草（Elymus nutans）是高寒地区建植和改良栽培草地的首选草种。虽然合理植株密度和氮素添加量是垂穗披碱草栽培草地稳产的关键因子,但两者之间是否存在最佳互作组合仍不清楚。采用盆栽试验,通过分析不同植株密度（58、102、146株/m²）和氮素添加量（0、200、400 mg/kg）组合状态下垂穗披碱草株高、单株分蘖数、地上生物量、地下生物量、根系体积和地上地下生物量比,以确定理论上是否存在植株密度和氮素添加量的最佳组合。结果表明：随植株密度增加,垂穗披碱草株高、地上生物量和地上地下生物量比值均先增加后降低,而单株分蘖数逐渐减小,根系体积和地下生物量先增加后保持相对稳定;随氮素添加量增加,垂穗披碱草单株分蘖数、地上生物量和地上地下生物量比值均表现为先增加后降低,地下生物量逐渐降低。植株密度与氮素添加量互作虽然对垂穗披碱草的根系体积和单株分蘖数没有显著影响,但两者互作显著影响了垂穗披碱草株高、地上生物量、地下生物量、地上地下生物量比（P<0.01）,这些指标与植株密度和氮素添加量的关系均表现为一个开口向下的抛物面。当植株密度为102株/m²和氮素添加量为200 mg/kg时,垂穗披碱草栽培草地产量最大,生物量分配最优。垂穗披碱草植株密度和氮素添加互作时理论上存在最佳组合,这为垂穗披碱草栽培草地的田间管理提供了理论依据。     

Belowground responses of <Emphasis Type="Italic">Picea asperata</Emphasis> seedlings to warming and nitrogen fertilization in the eastern Tibetan Plateau

The impacts of global climatic change on belowground ecological processes of terrestrial ecosystems are still not clear. We therefore conducted an experiment in the subalpine coniferous forest ecosystem of the eastern edges of the Tibetan Plateau to study roots of Picea asperata seedlings and rhizosphere soil responses to soil warming and nitrogen availability from April 2007 to December 2008. The seedlings were subjected to two levels of temperature (ambient; infrared heater warming) and two nitrogen levels (0 or 25 g m⁻²year⁻¹ N). We used a free air temperature increase from an overhead infrared heater to raise both air and soil temperature by 2.1 and 2.6°C, respectively. The results showed that warming alone significantly increased total biomass, coarse root biomass and fine root biomass of P. asperata seedlings. Both total biomass and fine root biomass were increased, but coarse root biomass was significantly decreased by nitrogen fertilization and warming combined with nitrogen fertilization. Warming induced a prominent increase in soil organic carbon (SOC) and NO₃ ⁻-N of rhizosphere soil, while nitrogen fertilization significantly decreased SOC and NH₄ ⁺-N of rhizosphere soil. The warming, fertilization and warming × N fertilization interaction decreased soil microbial C significantly, but substantially increased soil microbial N. These results suggest that nitrogen deposition combined with warmer temperatures under future climatic change possibly will have no effect on fine root production of P. asperata seedlings, but could enhance the nitrification process of their rhizosphere soils in subalpine coniferous forests.     

Yield of nitrogen from minimally disturbed watersheds of the United States

Watersheds of the US Geological Survey's Hydrologic Benchmark Network program were used in estimating annual yield of total nitrogen and nitrogen fractions (ammonium, nitrate, dissolved organic N, particulate N) in relation to amount of runoff, elevation, and watershed area. Only watersheds minimally disturbed with respect to the nitrogen cycle were used in the analysis (mostly natural vegetation cover, no point sources of N, atmospheric deposition of inorganic N < 10 kg ha^–1 y^–1). Statistical analysis of the yields of total nitrogen and nitrogen fractions showed that elevation and watershed area bear no significant relationship to nitrogen yield for these watersheds. The yields of total nitrogen and nitrogen fractions are, however, strongly related to runoff (r ² = 0.91 for total N). Annual yield increases as runoff increases, but at a rate lower than runoff; annual discharge-weighted mean concentrations decline as annual runoff increases. Yields of total nitrogen and most nitrogen fractions bear a relationship to runoff that is nearly indistinguishable from a relationship that was documented previously for minimally disturbed watersheds of the American tropics. Overall, the results suggest strong interlatitudinal convergence of yields and percent fractionation for nitrogen in relation to runoff.     

Biomass production and nitrogen contents of the CAM plants Kalanchoe daigremontiana and K. tubiflora in cultures with different nitrogen and water supply

Summary Two CAM plant species (Kalanchoe daigremontiana and K. tubiflora) were cultivated in pure cultures with two different levels of nitrogen and water supply. A comparison of the plant dry weight showed that the productivity was severely reduced under low nitrogen or/and water conditions. Additionally the proportion of the different organs contributing to the total biomass, and thus the ratio of root/shoot dry weight shifted substantially. At the same time the production of leaf buds in ratio to leaf biomass was increased. Concentrations of organic nitrogen and nitrate in the shoots were drastically reduced under low nitrogen or/and water conditions, but organic nitrogen concentration in the roots remained relatively high. The leaf conductance over a day was investigated only for K. daigremontiana, and decreased with reduced water and nitrogen supply. The results indicate that CAM plants do not have the predicted high nitrogen use efficiency. Under environmental stress they change from biomass production towards provisions for life preservation.     

Linking larval nutrition to adult reproductive traits in the European corn borer Ostrinia nubilalis

Throughout an organism's lifetime, resources are strategically allocated to many different functions, including reproduction. Reproduction can be costly for both sexes; females produce nutrient‐rich eggs, whereas males of many species produce large and complex ejaculates. In capital breeding insects, nutrients are mainly acquired during the larval period, yet allocation decisions impact the reproductive fitness of adults. The present study examines the effect of larval dietary nitrogen on both male and female reproductive traits in the European corn borer moth Ostrinia nubilalis Hübner, whose adults do not feed and whose males transfer a large, nitrogen‐rich spermatophore. One day post‐eclosion, O. nubilalis adults reared on one of three different diets (3.0%, 1.6%, or 1.1% nitrogen) are mated and two experiments are undertaken: one to measure nitrogen and carbon content of male ejaculates, and the other to determine female fecundity and fertility. Although male larval diet does not alter the percentage nitrogen content of adult somatic tissue, males reared on the higher nitrogen diet (3.0%) produce spermatophores with increased nitrogen relative to somatic nitrogen. Furthermore, females raised on the 3.0% nitrogen diet receive spermatophores with lower carbon : nitrogen ratios and thus more nitrogen. Overall, females lay more eggs as their larval dietary nitrogen increases, although they lay fewer eggs when their mates are raised on the higher (3.0%) nitrogen diet. This suggests that O. nubilalis females may use male‐derived nitrogen not to supplement egg production, but rather for somatic maintenance. Overall, the present study furthers our understanding of how larval diet can affect adult fitness in Lepidoptera.     

Costs of reproduction in Nyssa sylvatica: sexual dimorphism in reproductive frequency and nutrient flux

Summary We examined the influence of differential reproductive frequency between the sexes on tertiary (phenotypic) sex ratios in the the dioecious tree Nyssa sylvatica (Nyssaceae). Reproduction was evaluated in relation to sex, size and canopy exposure using flowering data collected from 1229 marked trees over a four year period. For subsets of each population we used data on flower number, fruit crop size, fruit/flower ratios, and individual flower and fruit mass to compare biomass invested in reproductive structures of males and females. We also examined seasonal changes in stem nitrogen and soluble carbohydrate content in relation to flower and fruit production for trees of each sex. Our results indicate that: 1) Male-biased tertiary sex ratios could be explained by more frequent reproduction by male trees; 2) Estimated secondary sex ratios based on sums of all known males and females were not significantly different from 1:1; 3) Flowering frequency of males and females was significantly related to plant size (DBH) and exposure of the canopy to light; 4) Estimtes of reproductive biomass allocation ranged from 1.36 to 10.8 times greater for females relative to males; 5) Flower production was related to stem nutrient status for both sexes, but nutrient depletion and its effect on subsequent flowering was much more pronounced for female trees. We conclude that less frequent flowering by female trees may result from depletion of stored reserves, and that differential flowering frequency in N. sylvatica may ultimately reduce apparent sexual differences in the costs of reproduction.     

荒漠草原植物群落结构及其稳定性对增水和增氮的响应

通过在荒漠草原开展增水和增氮野外控制试验,研究增水和增氮对荒漠草原植物群落结构、物种多样性及群落稳定性的影响。结果表明：（1）增水和增氮处理显著影响了荒漠草原植物群落结构和地上生物量,而对植物群落稳定性影响不显著（P>0.05）。增水处理显著增加了豆科和禾本科植物地上生物量（101.3%和57.9%）（P<0.05）;增水+增氮处理显著增加了植物群落盖度（43.2%）和地上生物量（112.4%）及不同功能群（禾本科和杂类草）植物盖度（75.5%和47.3%）和地上生物量（139.3%和85.7%）（P<0.05）。与增氮处理相比,增水+增氮处理显著增加了植物群落和不同功能群（禾本科和杂类草）植物高度、盖度和地上生物量（P<0.05）。（2）增水、增氮和增水+增氮处理均显著降低了植物群落Pielou指数（11.7%、8.7%和10.2%）（P<0.05）。（3）增水和增水+增氮处理提高了荒漠草原植物群落稳定性,而增氮处理降低了荒漠草原植物群落稳定性。增水处理荒漠草原植物群落稳定性效应大于增水+增氮处理。研究表明,荒漠草原植物群落结构受到氮沉降和降水增加的共同影响。增加降水对荒漠草原植物群落稳定性的积极效应可能会抵消部分氮沉降的消极影响,荒漠草原植物群落地上生物量及群落稳定性可能有所增加。     

A review of the life history,reproduction and phenology of Gracilaria

The basic life history of the red alga Gracilaria is of the three-phase Polysiphonia type but a number of species show deviations. Plants can bear both gametangia and tetrasporangia, either on separate parts of the thallus or on the same. Explanations include the in situ germination of tetraspores (allowing gametophytic thalli to be epiphytic on tetrasporophytes), the coalescence of spores or developing discs (resulting in chimaeras), mitotic recombination during cell division in the mature diploid thallus (resulting in patches of diploid male and female cells on the tetrasporophyte), a mutation eliminating the repression of female expression allowing haploid male plants to be bisexual and initial failure of cell walls to form during the development of tetraspores. Polyploids can be produced from plants with diploid gametangia. The sexes and phases are usually morphologically identical but gametophytes or their parts may be smaller. The growth rates of the sexes may differ and diploid juveniles may survive better than haploid. Neither polyploidy nor hybridization results in superior growth. The sex ratio is probably 1:1 but females may appear to be more abundant. Diploid and haploid phases are usually either about equal or diploids predominate, often depending on the type of substratum. At high latitudes reproduction peaks in late summer whereas in the tropics it may be high all year. In temperate regions growth rate is fastest and biomass highest in late summer; in the tropics peak biomass is mainly in the winter. Spermatia are effective for only a few h. Spores vary in size around 25 \m, diploid ones usually being larger. Cystocarps or tetrasporangia in the field may not currently be releasing spores. In the laboratory spore release shows a diurnal rhythm, peaking during the night or day according to the species. All the above attributes are potentially important in planning and executing Gracilaria cultivation.     

Selection of maternity dens by female polar bears in western Hudson Bay,Canada and the effects of human disturbance

We deployed 19 satellite radio collars on adult female polar bears (Ursus maritimus) in western Hudson Bay to study how pregnant females select maternity-den sites and the degree to which human disturbance might influence this process. To eliminate the possibility that being handled while pregnant might influence the final choice of a suitable site, the collars were attached 1 year before the females were expected to select dens. Our results indicated that pregnant females moved inland and selected an area to den shortly after coming ashore in the summer and remained there if not disturbed. We analyzed 24 years of consecutive data (1979–2002) to examine whether disturbance of pregnant females in the denning area in the autumn affected litter size or mass of cubs the following spring. We defined three levels of disturbance (none—bear not seen; low—bear seen but not handled; high—bear seen and handled), and found that disturbance had no effect on either litter size or the mass of male cubs. However, we did find that females handled in the autumn had significantly lighter female cubs than females that were not seen. Our disturbance of pregnant females occurred in August and September, which is only a few months prior to parturition; it is not known what the effects of disturbance might be in late October or early November, closer to the time when cubs are born.     

EFFECTS OF SALINITY,NITROGEN, AND POPULATION DENSITY ON THE SURVIVAL,GROWTH, AND REPRODUCTION OF ATRIPLEX TRIANGULARIS (CHENOPODIACEAE)

Populations of Atriplex triangularis were grown under laboratory conditions in a growth chamber and manipulated in an inland Ohio saline pond in order to examine the relative effects of salinity, nitrogen fertilization, and population density on growth, reproduction, and survival. For laboratory plants, nitrogen fertilization was the most important variable, with biomass and reproductive effort being greatest at the high nitrogen level. As salinity increased, biomass decreased only in plants not limited by nitrogen. Increasing density caused biomass per plant to decrease at both high and low nitrogen levels. For field plants, density was the most important variable, with biomass per plant and survival both decreasing as density increased. As density increased, size inequality among individuals increased but biomass per unit area and individual reproductive effort remained relatively constant. Nitrogen fertilization slightly enhanced survival, but did not affect biomass. It is suggested that density-dependent processes may be significant even in relatively harsh physical environments.