How to test different density-dependent fecundity hypotheses in an increasing or stable population

1. We report on a simulation study of increasing and stable populations working under two different hypotheses of density dependence of fecundity: the habitat heterogeneity hypothesis (HHH) and the individual adjustment hypothesis (IAH). Our aim is to find critical differences between the two regulatory hypotheses in natural populations. 2. Populations under HHH show a strong negative relationship between fecundity and the coefficient of variation of fecundity. We also found a strong negative relationship between fecundity and skewness, demonstrating that, as fecundity decreases, the form of the distribution of brood sizes changes, being more left-skewed due to more territories failing to produce any offspring. 3. This strong relationship was found only in the simulations of populations under HHH; whether increasing or stable, and under different ratios of good: poor territories and different population sizes. In contrast, no relationship between mean fecundity and skewness was found among simulations under IAH. 4. Populations under IAH also showed a significant relationship between mean fecundity and the coefficient of variation of fecundity, but with a lower slope than in populations under HHH. 5. In conclusion, skewness was found to be an adequate critical test that showed significant and strong relationships with mean fecundity only in populations under HHH, whether increasing or stable. This test is useful for species with a discrete distribution of offspring with a small number of integer categories, including most of the bird and mammal species.     

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.     

植物群落物种共存机制:负密度制约假说

负密度制约假说主要描述由于资源竞争、有害生物侵害(比如病原微生物、食草动物捕食)等, 同种个体之间发生的相互损害行为; 它主要强调同种个体之间的相互作用, 解释自然群落物种共存的机理; 负密度制约机制主要在小尺度上降低群落内同种个体生长率, 同时提高个体死亡率, 从而为其他物种的生存提供空间和资源, 促进物种共存。目前负密度制约假说的检验研究主要侧重密度制约、距离制约、群落补偿效应等三个方面。最近, 研究者又探讨了近缘物种之间由于对相似资源的竞争所产生的负效应, 扩展了负密度制约假说, 进而提出异群保护假说和谱系多样性制约假说。负密度制约假说引起生态学家长久的探讨和关注, 世界范围内大尺度森林动态样地的建立, 又为探索不同尺度上密度制约效应的研究提供了条件。然而目前的研究仍然存在不足, 比如负密度制约假说的检验受到其他因素的干扰、区域研究不平衡等。因此, 生态学家们仍然怀疑负密度制约效应调节群落物种共存的重要性, 但是目前的研究还没有发现否定负密度制约假说的充分证据。     

Is there genetic variation in the response to competition intensity in juvenile brown trout Salmo trutta?

Effects of intraspecific competition intensities on the relative performance (growth and movement) of juvenile brown trout Salmo trutta originating from nine different families were tested in tank experiments and in semi-natural streams. Both growth and movement differed consistently among families, indicating genetic variation in these traits. There were no significant interaction effects, however, between the intensity of competition and family origin on performance in either of the two experimental systems. Thus, genetic variation in response to competition intensity appeared to be limited in the population from which the juveniles used in this experiment originate.     

Is the shape of the density-growth relationship for stream salmonids evidence for exploitative rather than interference competition?

1. Empirical studies show that average growth of stream-dwelling salmon and trout often declines with increasing density in a characteristic concave relationship. However, the mechanisms that generate negative density-growth relationships in populations in natural streams are not certain. 2. In a recent study, Imre, Grant & Cunjak (2005; Journal of Animal Ecology, 74, 508-516) argue that density-dependent growth due to exploitative competition for prey causes the negative density-growth relationships for stream salmonids. They argue that the concave shape of empirical density-growth relationships is consistent with a simple model of exploitative competition and not consistent with interference competition for space. 3. We use a simple model to show that competition for space can yield concave density-growth relationships consistent with the empirical pattern when individuals compete for foraging sites that vary spatially in quality and lower-quality sites predominate. Thus, the predictions of the exploitative competition and spatial competition models overlap. 4. The shape of the density-growth relationship does not differentiate between candidate mechanisms underlying density-dependent growth for stream salmonids. Our results highlight the general problem with determining the mechanism driving an ecological process from patterns in observational data within the context of linking population demographics to habitat structure and animal behaviour.     

Sexual reproduction and diversity: Connection between sexual selection and biological communities via population dynamics

Sexual reproduction is a mysterious phenomenon. Most animals and plants invest in sexual reproduction, even though it is more costly than asexual reproduction. Theoretical studies suggest that occasional or conditional use of sexual reproduction, involving facultative switching between sexual and asexual reproduction, is the optimal reproductive strategy. However, obligate sexual reproduction is common in nature. Recent studies suggest that the evolution of facultative sexual reproduction is prevented by males that coerce females into sexual fertilization; thus, sexual reproduction has the potential to enforce costs on a given species. Here, the effect of sex on biodiversity is explored by evaluating the reproductive costs arising from sex. Sex provides atypical selection pressure that favors traits that increase fertilization success, even at the expense of population growth rates, that is, sexual selection. The strength of sexual selection depends on the density of a given species. Sexual selection often causes strong negative effects on the population growth rates of species that occur at high density. Conversely, a species that reduces its density is released from this negative effect, and so increases its growth rate. Thus, this negative density-dependent effect on population growth that arises from sexual selection could be used to rescue endangered species from extinction, prevent the overgrowth of common species and promote the coexistence of competitive species. Recent publications on sexual reproduction provide several predictions related to the evolution of reproductive strategies, which is an important step toward integrating evolutionary dynamics, demographic dynamics and community dynamics.     

不同倍性冬小麦种内竞争能力的比较研究

通过3个不同倍性冬小麦材料(两倍体栽培一粒、四倍体栽培两粒、六倍体现代品种长武134),在不同水分条件下进行密度实验,研究了不同材料的株高、生物量累积和分蘖动态的变化,以及产量对密度变化的反应。结果表明随着群体的增大,不同倍性材料个体间竞争明显加剧,相互抑制作用增强,种群内部个体大小等级差异增大;在不同群体下各倍性材料的个体生长存在差异,表现为四倍体栽培两粒竞争能力两倍体栽培一粒六倍体现代品种长武134,且长武134受种群大小影响最为显著,但长武134产量累积的投入比例最高,产量最高,低竞争能力的个体更适合生产上的需求,是群体高产的基础。研究结果为旱地小麦的高产栽培和育种提供了理论基础。     

木林子保护区优势种翅柃种群结构与空间分布格局

该研究以湖北木林子保护区优势种翅柃(Eurya alata)为研究对象,编制静态生命表,绘制存活曲线、死亡率和消失率曲线;采用成对相关函数分析各龄级的空间分布特征,双变量成对相关函数分析不同生活史阶段间的空间关系,为阐明木林子保护区常绿落叶阔叶混交林不同构建规律在物种多样性维持机制中的作用提供理论参考。结果显示:(1)翅柃属增长型种群,存活曲线趋于Deevey-Ⅱ型,死亡率和消失率曲线呈单峰型。(2)翅柃Ⅰ～Ⅴ龄级呈现聚集分布且随径级增加而减弱,Ⅵ、Ⅶ龄级在较大尺度上随机分布。(3)在完全随机零模型下,各生活史阶段均呈空间正关联;在先决条件零模型下,成年树与幼树、小树小尺度上呈无关联和负关联,其余生活史阶段间呈正关联,较大尺度上无关联。研究表明,翅柃种群幼树个体丰富,更新状态良好,在自然演替过程中,生境过滤、扩散限制和密度制约均可能驱动种群的构建过程。     

Simulating density-dependent relationships in south Texas northern bobwhite populations

Density dependence influences northern bobwhite (Colinus virginianus) reproduction and overwinter mortality. However, the functional forms of these density-dependent relationships or the factors that influence them during the annual life cycle events of this bird are not clear. We used a systems analysis approach with a compartment model based on difference equations (Δt = 3 months) for bobwhites in South Texas to simulate population behavior using 16 different functional forms of density-dependent production and overwinter mortality. During the reproductive season, a weak linear density-dependent relationship resulted in the longest population persistence (up to 100.0 yr), whereas a reverse-sigmoid density-dependent relationship had the worst population persistence (2.5–3.5 yr). Regarding overwinter mortality, a sigmoid or weak linear density-dependent relationship and a weak linear or no density-dependent reproduction relationship had the longest population persistence (87.5–100.0 yr). Weak linear density-dependent reproduction with either sigmoid or weak linear overwinter mortality produced stable fall population trends. Our results indicated that density dependence may have a greater influence on overwinter survival of bobwhites than previously thought. Inclusion of density-dependent functional relationships that represent both density-dependent reproduction and overwinter mortality, were critical for our simulation model to function properly. Therefore, integrating density-dependent relationships for both reproductive and overwinter periods of the annual cycle of bobwhite life history events is essential for conducting realistic bobwhite population simulation analyses that can be used to test different management scenarios in an integrated and interdisciplinary manner. © 2012 The Wildlife Society.     

Assessing the impact of density dependence in field populations of Aedes aegypti

Although many laboratory studies of intra-specific competition have been conducted with Ae. aegypti, there have been few studies in natural environments and none that examined density dependence in natural containers at normal field densities. Additionally, current mathematical models that predict Ae. aegypti population dynamics lack empirically-based functions for density-dependence. We performed field experiments in Tapachula, Mexico, where dengue is a significant public health concern. Twenty-one containers with natural food and water that already contained larvae were collected from local houses. Each container was divided in half and the naturally occurring larvae were apportioned in a manner that resulted in one side of the container (high density) having four times the density of the second side (low density). Larvae were counted and pupae were removed daily. Once adults emerged, wing span was measured to estimate body size. Density had a significant impact on larval survival, adult body size, and the time taken to transition from 4(th) instar to pupation. Increased density decreased larval survival by 20% and decreased wing length by an average of 0.19 mm. These results provide a starting point for a better understanding of density dependence in field populations of Ae. aegypti.     

Experimental manipulation of breeding density and litter size: effects on reproductive success in the bank vole

1. Reproductive success of individual females may be determined by density-dependent effects, especially in species where territory provides the resources for a reproducing female and territory size is inversely density-dependent.
2. We manipulated simultaneously the reproductive effort (litter size manipulation: ± 0 and + 2 pups) and breeding density (low and high) of nursing female bank voles Clethrionomys glareolus in outdoor enclosures. We studied whether the reproductive success (number and quality of offspring) of individual females is density-dependent, and whether females can compensate for increased reproductive effort when not limited by saturated breeding density.
3. The females nursing their young in the low density weaned significantly more offspring than females in the high density, independent of litter manipulation.
4. Litter enlargements did not increase the number of weanlings per female, but offspring from enlarged litters had lower weight than control litters.
5. In the reduced density females increased the size of their home range, but litter manipulation had no significant effect on spacing behaviour of females. Increased home range size did not result in heavier weanlings.
6. Mother's failure to successfully wean any offspring was more common in the high density treatment, whereas litter manipulation or mother's weight did not affect weaning success.
7. We conclude that reproductive success of bank vole females is negatively density-dependent in terms of number, but not in the quality of weanlings.
8. The nursing effort of females (i.e. the ability to provide enough food for pups) seems not to be limited by density-dependent factors.     

A shift from exploitation to interference competition with increasing density affects population and community dynamics

Intraspecific competition influences population and community dynamics and occurs via two mechanisms. Exploitative competition is an indirect effect that occurs through use of a shared resource and depends on resource availability. Interference competition occurs by obstructing access to a resource and may not depend on resource availability. Our study tested whether the strength of interference competition changes with protozoa population density. We grew experimental microcosms of protozoa and bacteria under different combinations of protozoan density and basal resource availability. We then solved a dynamic predator–prey model for parameters of the functional response using population growth rates measured in our experiment. As population density increased, competition shifted from exploitation to interference, and competition was less dependent on resource levels. Surprisingly, the effect of resources was weakest when competition was the most intense. We found that at low population densities, competition was largely exploitative and resource availability had a large effect on population growth rates, but the effect of resources was much weaker at high densities. This shift in competitive mechanism could have implications for interspecific competition, trophic interactions, community diversity, and natural selection. We also tested whether this shift in the mechanism of competition with protozoa density affected the structure of the bacterial prey community. We found that both resources and protozoa density affected the structure of the bacterial prey community, suggesting that competitive mechanism may also affect trophic interactions.     

长白山次生杨桦林种群空间点格局及密度制约效应

以长白山5.2 hm²次生杨桦林样地为研究对象,利用空间点格局分析的双相关函数g（r）以及随机标签零模型和案例-对照设计法,探讨了树木种群空间格局及其密度制约效应。研究结果表明：生境异质性对不同生活型以及不同树种的成熟个体在大尺度范围上具有强烈影响。剔除生境异质性带来的影响后,在14个常见树种中有12个树种在小尺度上呈显著的空间聚集分布格局;随着空间尺度的增加聚集性分布树种数量急剧下降,在18 m尺度上聚集率下降到0。全部14个常见树种均呈现出显著的密度制约效应,表明密度制约是调节温带森林树木种群空间结构的主要作用机制。此外,密度制约与物种多度呈负相关,与亚林层和灌木层树种的同种聚集强度呈正相关。并且随着空间尺度增大受密度制约影响的树种百分比逐渐减小,14个常见树种中有11个在0-1 m处达到最大的密度制约强度。     

Determinants of species‐area relationships for marsh‐nesting birds

ABSTRACT To clarify the underlying causes of the species‐area relationship in marsh‐nesting birds, I studied eight freshwater tidal marshes of the Connecticut River that differed in area, degree of isolation, mudflat cover, water cover, tidal regime, and extent of individual plant communities. I measured these habitat variables on aerial infrared photos, and surveyed bird populations by mapping the distribution of all birds in marshes under 5 ha in area and establishing 50‐m radius plots in marshes over 5 ha. From surveys, I determined species richness, population densities, and total populations. Analysis revealed a positive relationship between species richness and area, but no correlation between area and habitat heterogeneity. Other habitat variables were poor predictors of species richness. The lack of a relationship between habitat and species richness appeared to be a consequence of most vegetation types present not being sufficiently distinct for birds to differentially associate with them. I also found no relationship between bird population density and area, suggesting that habitat quality in marshes did not improve with increasing size, and species evenness declined with increasing richness because greater richness was associated with the presence of more rare species. Larger marshes had more rare species, species with larger populations, and species with a minimum threshold area for occurrence. Thus, my results are consistent with theoretical predictions that larger populations are less prone to local extinction and, as individuals are added to a community, more rare species are present.     

Interactions between density,home range behaviors,and contact rates in the Channel Island fox (Urocyon littoralis)

Many of the mechanisms underlying density‐dependent regulation of populations, including contest competition and disease spread, depend on contact among neighboring animals. Understanding how variation in population density influences the frequency of contact among neighboring animals is therefore an important aspect to understanding the mechanisms underlying, and ecological consequences of, density‐dependent regulation. However, contact rates are difficult to measure in the field and may be influenced by density through multiple pathways. This study explored how local density affects contact rates among Channel Island foxes (Urocyon littoralis) through two pathways: changes in home range size and changes in home range overlap. We tracked 40 radio‐collared foxes at four sites on San Clemente Island, California. Fox densities at the four sites ranged from 2.8 ± 1.28 to 42.8 ± 9.43 foxes/km². Higher fox densities were correlated with smaller home ranges (R² = 0.526, F_1,38 = 42.19, P < 0.001). Thirty foxes wore collars that also contained proximity loggers, which recorded the time and duration of occasions when collared foxes were within 5 m of one another. Contact rates between neighboring fox dyads were positively correlated with home range overlap (R² = 0.341, P = 0.008), but not fox density (R² = 0.012, P = 0.976). Individuals at high densities had more collared neighbors with overlapping home ranges (R² = 0.123, P = 0.026) but not an increase in the amount of contact between individual neighbors. This study was the first time contact rates were directly measured and compared to density and home range overlap. Results suggest that foxes exhibit a threshold in their degree of tolerance for neighbors, overlap is a reliable index of the amount of direct contact between island foxes, and disease transmission rates will likely scale with fox density.