An experimental analysis of self-thinning in juvenile steelhead trout

Mobile animal populations have been proposed to decline in density according to a slope based on the allometry of metabolic requirements or space requirements. In salmonid fishes, metabolic rate and food consumption scale to body mass by the exponent 0.87 and 0.73, respectively; whereas the territory size of steelhead trout scales to body mass by the exponent 0.86. Experimental cohorts of juvenile steelhead trout ( Oncorhynchus mykiss ) were used to test the hypothesis that mobile animal populations composed of individuals with indeterminate growth decline in density as a result of self-thinning. After controlling for experimentally manipulated levels of food abundance and stocking density, cohorts of steelhead trout declined in density with increasing body size according to a slope closest to the allometry of food consumption. Densities of steelhead trout were inversely related to average mass by the exponent −0.74. Despite the similarity to the food consumption slope, a relatively wide confidence interval also precluded distinguishing the slope either the metabolic rate or territory size slopes. Data from the literature were also examined to determine if there was general support for the idea of self-thinning in natural populations of stream-dwelling salmonid fish. Although not all data suggest that populations of salmonids in streams decline as a result of density-dependent intraspecific competition, at least some appear to fit the idea of self-thinning; especially when density is above a minimum level of habitat saturation.     

Allometry of territory size and metabolic rate as predictors of self-thinning in young-of-the-year Atlantic salmon

1. Self-thinning is a progressive decline in population density caused by competitively induced losses in a cohort of growing individuals and can be depicted as: log₁₀ (density) = c − β log₁₀ (body mass).
2. In mobile animals, two mechanisms for self-thinning have been proposed: (i) the space hypothesis predicts that maximum population density for a given body size is the inverse of territory size, and hence, the self-thinning slope is the negative of the slope of the allometric territory-size relationship; (ii) the energetic equivalence hypothesis predicts that the self-thinning slope is the negative of the slope of the allometric metabolic rate relationship, assuming a constant supply of energy for the cohort.
3. Both hypotheses were tested by monitoring body size, population density, food availability and habitat for young-of-the-year Atlantic salmon ( Salmo salar ) in Catamaran Brook, New Brunswick. The results were consistent with the predictions of the space hypothesis. Observed densities did not exceed the maximum densities predicted and the observed self-thinning slope of −1·16 was not significantly different from the slope of −1·12, predicted by the allometry of territory size for the population under study.
4. The observed self-thinning slope was significantly steeper than −0·87, predicted by the allometry of metabolic rate, perhaps because of a gradual decline in food abundance over the study period. The decline in density was more rapid in very shallow sites and may have been partly caused by a seasonal change in water depth and an ontogenetic habitat shift rather than solely by competition for food or space.
5. The allometry of territory size may be a useful predictor of self-thinning in populations of mobile animals competing for food and space.     

Relationship between the virtual dynamic thinning line and the self-thinning boundary line in simulated plant populations

The self-thinning rule defines a straight upper boundary line on log-log scales for all possible combinations of mean individual biomass and density in plant populations. Recently, the traditional slope of the upper boundary line, -3/2, has been challenged by -4/3 which is deduced from some new mechanical theories, like the metabolic theory. More experimental or field studies should be carried out to identify the more accurate self-thinning exponent. But it＇s hard to obtain the accurate self-thinning exponent by fitting to data points directly because of the intrinsic problem of subjectivity in data selection. The virtual dynamic thinning line is derived from the competition-density （C-D） effect as the initial density tends to be positive infinity, avoiding the data selection process. The purpose of this study was to study the relationship between the virtual dynamic thinning line and the upper boundary line in simulated plant stands. Our research showed that the upper boundary line and the virtual dynamic thinning line were both straight lines on log-log scales. The slopes were almost the same value with only a very little difference of 0.059, and the intercept of the upper boundary line was a little larger than that of the virtual dynamic thinning line. As initial size and spatial distribution patterns became more uniform, the virtual dynamic thinning line was more similar to the upper boundary line. This implies that, given appropriate parameters, the virtual dynamic thinning line may be used as the upper boundary line in simulated plant stands.     

Interspecific habitat associations of juvenile salmonids in Lake Ontario tributaries: implications for Atlantic salmon restoration

Diel variation in habitat use of subyearling Chinook salmon (Oncorhynchus tshawytscha), subyearling coho salmon (O. kisutch), yearling steelhead (O. mykiss), and yearling Atlantic salmon (Salmo salar) was examined during the spring in two tributaries of Lake Ontario. A total of 1318 habitat observations were made on juvenile salmonids including 367 on steelhead, 351 on Chinook salmon, 333 on Atlantic salmon, and 261 on coho salmon. Steelhead exhibited the most diel variation in habitat use and Chinook the least. Juvenile salmonids were generally associated with more cover and larger substrate during the day in both streams. Interspecific differences in habitat use in both streams occurred with Atlantic salmon (fast velocities) and coho salmon (pools) using the least similar habitat. Chinook salmon and Atlantic salmon used similar habitat in both streams. These findings should help guide future management actions specific to habitat protection and restoration of Atlantic salmon in Lake Ontario tributaries.     

Density-dependent Mortality Induced by Low Nutrient Status of the Substrate

The hypothesis that changing the fertility level of the substratewould change the self-thinning line (different slope or intercept)followed by high-density populations was tested by sowing populationsof Ocimum basilicum L. at two densities on a soil-based pottingmix adjusted to three fertility levels (F0, F1 and F2). Fertilitylevel significantly affected the slope of the thinning linesfor both shoot and root biomass. For shoot biomass, more mortalityoccurred per unit increase in biomass as fertility level declined(the slope of the thinning line became flatter). The slope ofthe log shoot biomassvs. log density relationship was -0.5 atthe F2-, zero at the F1-, and 0.94 at the F0-fertility. Forthe log root biomassvs. log density lines, slopes were zeroat the F2- and F0-fertility levels, and -0.32 at F1. Packingof shoot biomass into canopies of individual plants correlatedwell with observed exponents of self-thinning lines at the F2-and F1-fertility level. Plants at the F2-fertility level requiredmore canopy space to support a given shoot biomass than plantsat F1, indicating that shoot competition was more intense atthe F2-fertility level for a given biomass. Leaf area indexand size inequality also increased with fertility level fora given shoot biomass. Density-dependent mortality in populationsgrown at the F0-fertility level was highly unusual in havinga positive slope for the shoot biomass vs. density relationship.Shoot growth per plant was static as density declined in theF0-populations; however, root growth per plant increased. Allmeasurements of shoot growth (mass, height, canopy extension,leaf area) remained static in the F0-populations: root massand length increased in comparison. It is argued that root competitionbecame sufficiently intense to cause the density-dependent mortalityseen at the F0-fertility level, with little contribution ofshoot competition to mortality. Copyright 1999 Annals of BotanyCompany Ocimum basilicum, self-thinning, root competition, shoot competition, fertility level and competition, density-dependent mortality, allometric self-thinning.     

Density dependence across multiple life stages in a temperate old-growth forest of northeast China

Recent studies on species coexistence suggest that density dependence is an important mechanism regulating plant populations. However, there have been few studies of density dependence conducted for more than one life-history stage or that control for habitat heterogeneity, which may influence spatial patterns of survival and mask density dependence. We explored the prevalence of density dependence across multiple life stages, and the effects of controlling for habitat heterogeneity, in a temperate forest in northeast China. We used generalized linear mixed-effects models to test for density-dependent mortality of seedlings and spatial point pattern analysis to detect density dependence for sapling-to-juvenile transitions. Conspecific neighbors had a negative effect on survival of plants in both life stages. At the seedling stage, we found a negative effect of conspecific seedling neighbors on survival when analyzing all species combined. However, in species-level analyses, only 2 of 11 focal species were negatively impacted by conspecific neighbors, indicating wide variation among species in the strength of density dependence. Controlling for habitat heterogeneity did not alter our findings of density dependence at the seedling stage. For the sapling-to-juvenile transition stage, 11 of 15 focal species showed patterns of local scale (<10 m) conspecific thinning, consistent with negative density dependence. The results varied depending on whether we controlled for habitat heterogeneity, indicating that a failure to account for habitat heterogeneity can obscure patterns of density dependence. We conclude that density dependence may promote tree species coexistence by acting across multiple life-history stages in this temperate forest.     

Geomorphology and fish assemblages in a Piedmont river basin, U.S.A.

1. We investigated linkages between fishes and fluvial geomorphology in 31 wadeable streams in the Etowah River basin in northern Georgia, U.S.A. Streams were stratified into three catchment sizes of approximately 15, 50 and 100 km², and fishes and geomorphology were sampled at the reach scale (i.e. 20–40 times stream width). 2. Non‐metric multidimensional scaling (NMDS) identified 85% of the among‐site variation in fish assemblage structure and identified strong patterns in species composition across sites. Assemblages shifted from domination by centrarchids, and other pool species that spawn in fine sediments and have generalised food preferences, to darter‐cyprinid‐redhorse sucker complexes that inhabit riffles and runs, feed primarily on invertebrates, and spawn on coarser stream beds. 3. Richness and density were correlated with basin area, a measure of stream size, but species composition was best predicted (i.e. |r| between 0.60–0.82) by reach‐level geomorphic variables (stream slope, bed texture, bed mobility and tractive force) that were unrelated to stream size. Stream slope was the dominant factor controlling stream habitat. Low slope streams had smaller bed particles, more fines in riffles, lower tractive force and greater bed mobility compared with high slope streams. 4. Our results contrast with the ‘River Continuum Concept’ which argues that stream assemblages vary predictably along stream size gradients. Our findings support the ‘Process Domains Concept’, which argues that local‐scale geomorphic processes determine the stream habitat and disturbance regimes that influence stream communities.     

Density dependence is prevalent in a heterogeneous subtropical forest

Although negative conspecific density dependence among neighbours is widely studied, the general prevalence of the effects is still poorly understood due to a lack of studies from zonal forests other than the tropics. In addition, the detection of density dependence may be confounded by the influence of habitat heterogeneity. Here we examined the spatial distributions of 47 common tree species (diameter at breast height≥1  cm) using the pair-correlation function g(r) in a fully mapped 24-ha subtropical forest in China. We first investigated whether habitat heterogeneity influenced tree distributions, and then examined the conspecific tree patterns and density dependence after removing the effects of habitat heterogeneity. We found that the forest plot exhibited strong large-scale heterogeneity in the distribution of both large adult trees of different growth forms and individual species. After the habitat heterogeneity was accounted for, 39 of the 47 species (83.0%) were found to exhibit density dependence predominantly at close distances among neighbors. Our findings highlight density dependence as a prevalent mechanism for regulating the population spatial structure of most tree species in the species-rich subtropical forest studied here. Furthermore, the occurrence of density dependence is closely associated with species abundance and the strength of conspecific aggregation at local scales. Abundant species with high strength of conspecific aggregation tend to show density dependence.     

贺兰山马鹿冬季取食和卧息生境选择

2007 年12 月至2008 年1 月,在贺兰山地区,利用痕迹检验法和直接观察法对马鹿阿拉善亚种的冬季取食和卧息生境选择进行研究。通过在选定的15 条沟段里进行调查,共测定了72 个取食利用样方,59 个卧息利用样方和131 个对照样方的18 种生态因子。结果表明,马鹿在冬季偏好的取食地为平滑起伏坡,以酸枣、柳为优势乔木、混合型树林或空地,乔木高大稀疏、间距较远,灌木密度大,草本盖度较高,位于< 15°的半阴半阳坡的下坡位,距裸岩远,隐蔽度高;偏好的卧息地在山地疏林草原带的平滑起伏坡上,以酸枣、柳为优势乔木、混合型树林或空地,乔木高大稀疏,灌木矮小、稀疏且距离较远,草本盖度较高,位于< 15°阳坡的下坡位,远离裸岩,隐蔽程度高。马鹿冬季的取食和卧息生境在草本盖度和隐蔽度上差异极显著。相对于卧息生境,马鹿冬季的取食生境对草本盖度和隐蔽度要求更低一些。马鹿冬季取食地的资源选择函数为1.155 - 0.149 × 乔木高度- 0.066 × 草本盖度+ 0.190 × 坡度,模型的正确判别率为86.8% ;马鹿冬季卧息地的资源选择函数为- 30.936
+ 0.494 × 乔木高度+ 0.257 × 坡度-0.002 × 海拔高度+ 0.387 × 隐蔽度,模型的正确判别率为95.8% 。食物、隐蔽条件以及贺兰山的独特地形特征是影响马鹿冬季取食和卧息生境选择的主要因素。     

Mortality and Self-thinning in Postfire Black Spruce

Using age-structure determinations on both living and dead stemsin censused plots, coupled with stem analysis techniques, anhistorical picture of mortality and above-ground tree stem growthwas recreated for ten stands dominated by black spruce in northeasternOntario, Canada. No evidence of mortality was seen in any plot prior to 30 yearsfollowing postfire initiation. Each of the eight oldest standsshowed a linear decline in numbers for a 20–25 year period.The steepness of the mortality slope was proportional to initiallive stem density within and among plots during this phase.The final 10–20 years was marked by a less steep declinein numbers. The log density vs log mean tree volume curves in the eightoldest stands were doubly asymptotic and were fitted to a logisticcurve very tightly in each case. At the point of inflectionthe curves' slopes ranged from –2.14 to –3.89. However,log density vs log mean stem volume among stands at this pointof inflection had a slope of –0.96. Reasons for the inconsistency between within-stand and among-standself-thinning estimates are considered, as well as the poorfit to the –3/2 rule. Ecosystem processes related to thechange in nutrient relations during stand growth are identifiedas a prime influence on self-thinning behaviour in natural blackspruce stands. Mortality, stem analysis, self-thinning, Picea mariana, black spruce     

Effect of Localized Placement of Nutrients on Root Competition in Self-thinning Populations

The hypothesis that increased root competition can lower theslope and/or intercept of the self-thinning line traversed byplant populations was tested using localized placement of nutrientsto increase root competition. Localized placement of nutrientswill result in increased root competition, if average inter-rootdistances are reduced, and if nutrients are in limiting supply.It was predicted that high-density, nutrient-limited populationsof Ocimum basilicum L. grown with localized placement of nutrientswould self-thin along a lower biomass–density line thannon-localized controls. This was tested at two fertility levelson a soil-based potting medium in expt 1, and at one fertilitylevel on washed sand in expt 2. Localized placement of nutrients significantly reduced the elevation(intercept) of the self-thinning line for both shoot and rootbiomass in expt 2. In expt 1, at the higher-fertility level,localized placement of nutrients had no significant effect;at the lower fertility level, localization had no significanteffect on thinning lines for shoot biomass, and resulted ina zero slope of the thinning line for root biomass. Canopy-based models of self-thinning failed to account for thereduction in the thinning-line intercept observed in expt 2.In both experiments, localized placement of nutrients resultedin a higher proportion of total root length being located inthe localization zone, which would result in a reduction inthe average inter-root distance. This would intensify root competitionunder conditions of nutrient limitation. The hypothesis thatintensified root competition would lower the self-thinning lineis supported by the results of expt 2. Localized placement of nutrients; root competition; shoot competition; root–shoot allocation; self-thinning; Ocimum basilicum ; sweet basil     

Body size–density relationship for Mytilus edulis in an experimental food-regulated situation

We grew mussels ( Mytilus edulis ) under two different food regimes and eight population density levels to estimate the joint effects of density and biomass on their growth and survival and to determine the shape of the biomass–density ( B–N) relationship. Mussels were reared for 22 months, between December 1994 and October 1996, in 1-L experimental chambers supplied with natural seston. Growth in shell length, individual wet mass and ash free dry mass ( m ) decreased with decreasing food availability and increasing population density. Survival was negatively correlated with density but did not differ significantly between food regimes during the first year. Variations in concentration of available food did not alter the effects of crowding on mussels, as judged from the slopes of the body size–density curves. The general patterns exhibited by B–N curves were not consistent with expectations since we found 1) no classical competition–density (C–D) effect as reported in plants at intermediate competition levels, and 2) a slope of −0.648 for m–N curves in both food regimes, which was higher than expected from self-thinning (ST) theory. This value does not support present food-driven ST theory. This study introduces an unusual m–N region which combines properties of both ST and C–D effect.     

Habitat diversity and species diversity: testing the competition hypothesis with juvenile salmonids

I experimentally tested two predictions of the hypothesis that the positive relationship between habitat diversity and species diversity arises from a reduction in the negative effects of interspecific competition. By allowing species to partition habitat and avoid competition, habitat diversity should 1) facilitate the addition of an ecologically intermediate species into an existing community, and 2) reduce the negative effects of that species on existing members of the community. I tested these predictions with juveniles of three sympatric salmonid species: coho salmon ( Oncorhynchus kisutch ), steelhead trout ( O. mykiss ), and cutthroat trout ( O. clarki ), which in natural streams occupy deep low-velocity pools, shallow high-velocity riffles, and intermediate habitats, respectively. I introduced two (coho and steelhead) and three species communities into each of three artificial stream habitats: pools, riffles, and diverse. The results provide partial support for the predictions. Cutthroat trout grew fastest in the diverse stream habitat. Though habitat diversity did not eliminate the negative effects of competition, in the three species community coho and steelhead grew as fast in the diverse habitat as in either homogeneous habitat. The results are consistent with data on species number and evenness from natural communities, where variation along other niche axes confounds the relationship between habitat diversity, interspecific competition, and species diversity.     

Self-thinning lines differ with fertility level

The effect of variations in fertility level of the substrate on the self-thinning lines followed by populations of Ocimum basilicum L. was investigated experimentally by establishing populations over a range of densities at two fertility levels. Populations from each fertility level followed different self-thinning lines for shoot biomass. Self-thinning began at a lower biomass in populations grown at the higher fertility level; the subsequent slope of the thinning line was –0.5 for these stands on a log shoot biomass versus log density plot. The slope of the self-thinning line was flatter (–0.29) at the lower fertility level. Fitting the self-thinning line by the Structural Relationship rather than the Major Axis made little difference to line estimates. Biomass packing differed with fertility level, with plants from the higher fertility stands requiring more canopy volume for given shoot biomass than plants from lower fertility levels. Biologically, this would mean shoot competition intensified more rapidly at the higher fertility level as biomass accumulated in stands. The difference in slope between fertility levels was associated with changes above- and belowground. The radial extension of the canopy versus shoot mass relationships of individual plants differed with fertility level. Plants at the lower fertility level allocated more biomass to root growth, and had less leaf area per unit root length. The differences in slope of the self-thinning lines may have been because of differences in the radial extension of the canopy versus shoot mass relationships of individual plants at each fertility level, and/or to an increase in root competition at the lower fertility level.     

Two‐phase self‐thinning in stream‐living juveniles of lake‐migratory brown trout Salmo trutta L. Compatibility between linear and non‐linear patterns across populations?

The self-thinning rule establishes that, in resource limited populations, increased per-capita requirements by individuals with indeterminate growth may limit density. This invariant rule relating log-log linear declines of cohort density with increased body mass has been well established in sessile organisms. However, few studies have assessed the occurrence of self-thinning in mobile organisms and still fewer have demonstrated its operation in natural situations. In Bisballe Baeck, where brown trout individuals experienced the simultaneous operation of density-dependence on growth and mortality, increased body mass explained 89.0–98.4% of the variations in density. However, relative to the invariant linear line, the decline in density with increased body mass described distinctly nonlinear, two-phase trajectories. An initial phase of variable slopes continued until body mass attained ≈10.2 g, at which the direction of the trajectory switched to a second phase of substantially steeper slopes or severe declines of density per mass unit increment. As inferred by the highly significant relationships between these two sets of slopes and annual recruitment, the two phases were consistent with a major assumption of self-thinning: the rate at which density declined with increased body mass increased with the intensity of intraspecific competition. The results of this study provide prima facie evidence for the occurrence of self-thinning in a mobile species based on observational data. A further comparison among density–body mass relationships described for resident, anadromous and lake-migratory brown trout suggests consistent two-phase self-thinning patterns across populations.     

冬季清凉峰山区小麂和野猪的生境选择及差异

从2005 年11 月5 日至2006 年1 月21 日,为了评估小麂和野猪的生境选择及其差异,我们在浙江清凉峰国家级自然保护区核心区共设置了248 个样方群。研究发现,两者都选择平缓坡的灌丛植被、草本密度小、离住宅较近和离隐蔽物距离适中的生境,表现了它们的生境选择有一定的重叠性。但是它们也表现出生境选择的一些差异,小麂选择乔木密度适中、灌丛密度大和郁闭度适中的生境,而野猪选择乔木密度小的生境,随机地使用各类灌丛密度和郁闭度的生境。小麂还选择北坡、离水源距离较远的生境,野猪只是随机地使用不同的坡向和离水源距离的生境。分析两者共存的机制,我们发现小麂采取隐藏策略而野猪采取逃跑策略来避敌和利用资源,即它们之间存在与反捕食策略相关的生境选择差异。
     

Plant diversity in four chalk grassland sites with different aspects

Plant diversity and its components was studied on four chalk grassland sites in southeast England over a fourteen-month period. The sites were almost adjacent, and each faced one of the major compass points, N, S, E and W. Aboveground vegetation was recorded at monthly intervals by cover repetition, using a point quadrat technique. Seasonal patterns in the development of leaf area, bare ground, number of species present and diversity were similar on all sites. The N and W facing slopes and the S and E facing slopes resembled each other in several characteristics, while these two pairs of slopes differed markedly from each other. Thus, (i) there were more grasses but fewer forbs on the N and W slopes, (ii) overall diversity and forb diversity were lower on the N and W slopes but grass diversity was higher, (iii) equitability was lower on the N and W slopes, (iv) percentage similarity of begetation present was high when the N slope was compared with the W and when the S slope was compared with the E; other comparisons gave lower percentage similarity. It is proposed that the greater abundance of Carex flacca on the N and W slopes is important in causing these differences, and that it results in lower diversity on these slopes by occupying part of the habitat of the species-rich forb community which occurs on the S and E slopes. Dominance-diversity curves range from lognormal to geometric in form; some interpretation of changes they display can be made by assuming that the community is in a non-equilibrium state caused by seasonal environmental fluctuations each winter.     

Environmental factors influencing the regional distribution and local density of a small benthic fish: the stoneloach (<Emphasis Type="Italic">Barbatula barbatula</Emphasis>)

We investigated the relationships between different environmental variables and the spatial distribution patterns of the stoneloach (Barbatula barbatula) at the stream system, the stream site, and the mesohabitat (riffle/pool) scales in south-western France. Stoneloach occurred at 240 sites (out of 554 sampling sites), chiefly close to the source, in areas at low elevation and with weak slopes. Population density at a site was primarily influenced by physical conditions. Stream width was positively related to the probability of presence of stoneloach within the stream system, but negatively related to local density. These results indicate that stoneloaches can occur in a wide range of streams, but they are less abundant in wide rivers, probably because of lower habitat heterogeneity. Slope was negatively correlated to both fish presence at the regional scale and local density, suggesting that stoneloach’s swimming performance were weak under greater erosive forces. These results suggested that the distribution of populations and the density of stoneloach were governed by the suitability of physical habitat. Multi-scale studies of factors influencing a species’distribution allow to integrate patterns observed at different scales, and enhance our understanding of interactions between animals and their environment. The use of few pertinent variables in successful final models could reduce the effort and cost of data collection for water management applications.     

Fine‐scale habitat heterogeneity influences browsing damage by elephant and giraffe

Effects of large mammalian herbivores on woody vegetation tend to be heterogeneous in space and time, but the factors that drive such heterogeneity are poorly understood. We examined the influence of fine‐scale habitat heterogeneity on the distribution and browsing effects of two of the largest African terrestrial mammals, the elephant and giraffe. We conducted this study within a 120‐ha (500 x 2,400 m) ForestGEO long‐term vegetation monitoring plot located at Mpala Research Center, Kenya. The plot traverses three distinct topographic habitats (“plateau,” “steep slopes,” and “valley”) with contrasting elevation, slope, soil properties, and vegetation composition. To quantify browsing damage, we focused on Acacia mellifera, a palatable tree species that occurs across the three habitat categories. Overall tree density, species richness, and diversity were highest on the steep slopes and lowest on the plateau. Acacia mellifera trees were tallest and had the lowest number of stems per tree on the steep slopes. Both elephant and giraffe avoided the steep slopes, and their activity was higher during the wet season than during the dry season. Browsing damage on Acacia mellifera was lowest on the steep slopes. Elephant browsing damage was highest in the valley, whereas giraffe browsing damage was highest on the plateau. Our findings suggest that fine‐scale habitat heterogeneity is an important factor in predicting the distribution of large herbivores and their effects on vegetation and may interact with other drivers such as edaphic variations to influence local variation in vegetation structure and composition.