The effects of landscape structure on space competition and alternative stable states

Many species that compete for space live on heterogeneous landscapes and interact at local scales. The quality, amount, and structure of landscapes may have considerable impact on the ability of species to compete or coexist, yet basic models of space competition do not include that level of detail. We model space competition between two species with positive feedback through recruitment facilitation, which creates the potential for alternative stable states to occur. We compare the predictions of a spatially implicit model with a simulation model that includes explicit space and landscape structure. We create structured landscapes in which we specify the amount of habitat and degree of fragmentation and ask how landscape structure, dispersal strategy, and scale affect the presence of alternative stable states, or bistability. We find that structured landscapes can reduce the range of parameter values that lead to bistability in our model, but they do not eliminate bistability. The type of landscape and the dispersal distance for each species also influence the amount of environmental change needed for abrupt community shifts to occur. Coexistence of the two competitors is possible under certain conditions when connectivity is low. Consequently, landscape structure may lead to considerable disparity between the predictions of simple models and actual dynamics on complex landscapes during environmental change.     

基于景观连接度的斑块分级的尺度效应

认识到尺度对景观格局分析结果的影响,生态学家们越来越多地进行尺度效应的相关研究.已有的尺度效应研究多注重描述景脱格局指数对尺度变化的响应,这只能对景观的状态进行描述.无法辨别那些对景观功能具有重要意义的关键景观元素.文章选取8个景观连接度指数.通过改变最小制图单元和物种的搜索范围,研究尺度对斑块分等定级的影响,并据此确定各连接度指数的尺度敏感性.结果表明:景观中斑块的相对重要性随尺度变化而变化,其变化程度因指数而异.其中,AWF为尺度效应最不明显的指数,PC和IIC次之,其余指数(NL,NC,LCP,H和F)的尺度效应均十分显著;在衡量景观的连通性时,除IIC外的其他所有指数的尺度效应均十分显著.由此说明,根据连接度指数确定的斑块相对重要性的尺度敏感性与该指数本身的尺度敏感性无关.     

1995至2004年广州土地利用格局变化与驱动分析

综合应用景观生态学、统计学基本原理,以及3S技术、手段,以广州市为研究对象,基于TM和SPOT影像数据,运用ERDAS、ARC/INFO、FRAGSTATS和SAS等GIS和统计学工具平台,分析了广州市1995到2004景观格局状况及其动态变化.研究结果表明:近10年间,广州市的耕地、林地和水域等自然、半自然景观面积减少,其中耕地面积减少的最多,而居民工矿用地和交通路网用地等人文景观面积都在增加,园地和湿地等半自然景观面积也在增加;斑块数量和边缘密度呈增加之势,景观破碎化程度升高;景观多样性指数的升高和景观聚集度的下降,景观异质性增加;景观形状指数增加,景观形状复杂化;影响景观格局变化的驱动因子主要是建筑密度、道路密度、人口密度和第三产业结构比例.根据因子分析的结果,从保护农地、降低景观破碎化程度等方面提出了城市景观的优化建议.     

The effects of seston food quality on planktonic food web patterns

In planktonic food webs, the conversion rate of plant material to herbivore biomass is determined by a variety of factors such as seston biochemical/elemental composition, phytoplankton cell morphology, and colony architecture. Despite the overwhelming heterogeneity characterizing the plant–animal interface, plankton population models usually misrepresent the food quality constraints imposed on zooplankton growth. In this study, we reformulate the zooplankton grazing term to include seston food quality effects on zooplankton assimilation efficiency and examine its ramifications on system stability. Using different phytoplankton parameterizations with regards to growth strategies, light requirements, sinking rates, and food quality, we examined the dynamics induced in planktonic systems under varying zooplankton mortality/fish predation, light conditions, nutrient availability, and detritus food quality levels. In general, our analysis suggests that high food quality tends to stabilize the planktonic systems, whereas unforced oscillations (limit cycles) emerge with lower seston food quality. For a given phytoplankton specification and resource availability, the amplitude of the plankton oscillations is primarily modulated from zooplankton mortality and secondarily from the nutritional quality of the alternative food source (i.e., detritus). When the phytoplankton community is parameterized as a cyanobacterium-like species, conditions of high nutrient availability combined with high zooplankton mortality led to phytoplankton biomass accumulation, whereas a diatom-like parameterization resulted in relatively low phytoplankton to zooplankton biomass ratios highlighting the notion that high phytoplankton food quality allows the zooplankton community to sustain relatively high biomass and to suppress phytoplankton biomass to low levels. During nutrient and light enrichment conditions, both phytoplankton and detritus food quality determine the extent of the limit cycle region, whereas high algal food quality increases system resilience by shifting the oscillatory region towards lower light attenuation levels. Detritus food quality seems to regulate the amplitude of the dynamic oscillations following enrichment, when algal food quality is low. These results highlight the profitability of the alternative food sources for the grazer as an important predictor for the dynamic behavior of primary producer–grazer interactions in nature.     

The effects of opportunistic and intentional predators on the herding behavior of prey

In this article, we study how predator behavior influences the aggregation of prey into herds. Game-theoretic models of herd formation are developed based on different survival probabilities of solitary prey and prey that join the herd and on the predator's preference of what type of prey to search for. For an intentional predator that will only pursue its preferred type of prey, a single herd with no solitaries cannot emerge unless the herd acts as a prey refuge. If neither prey choice provides a refuge, it is shown that an equilibrium always exists where there are both types of prey and the predator does not always search for the same type of prey (i.e., a mixed equilibrium exists). On the other hand, if the predator is opportunistic in that it sometimes shifts to pursue the type of prey that is observed first, there may be a single herd equilibrium that does not act as a prey refuge when there is a high level of opportunistic behavior. For low opportunistic levels, a mixed equilibrium is again the only outcome. The evolutionary stability of each equilibrium is tested to see if it predicts the eventual herding behavior of prey in its corresponding model. Our analysis confirms that both predator and prey preferences (for herd or solitary) have strong effects on why prey aggregate. In particular, in our models, only the opportunistic predator can maintain all prey in a single herd that is under predation risk.     

中国能源利用效率区域差异基尼系数分析

基尼系数是经济学中对贫富差距量化评价的重要工具.文章从基尼系数的经济学内涵出发,将基尼系数的概念和意义应用于地区能源利用效率分析.以全国各地区的能源消耗分配为例,应用基尼系数法分析了地区总体经济发展水平,地区工业化程度,地区经济重型化程度对能源消费强度分配公平性的影响,并提出了以绿色贡献系数作为判断不公平因子的依据.绘制各影响因素的洛伦茨曲线,计算出中国2006年的能源消耗基尼系数,根据基尼系数和绿色贡献系数对能源消费强度地区差异进行量化分析,最后得出合理的结论.     

Fire-climate history and landscape patterns of high burn severity areas on the California southern and central coast

There is mounting evidence that fire size and severity have been growing on the central and southern California coastal landscape over the past several decades. Landsat satellite data was analyzed for the 20 largest fires on the Central California coast since 1984 to determine the relationships between climate/weather conditions at the time of ignition and the size of high burn severity (HBS) areas. The study also examined the relationship between area burned and landscape patterns of HBS coverage, including patch size, edge complexity, perimeter-to-area ratio, and aggregation metrics. Results showed that climate conditions at the time of ignitions have been significant controllers of the total area of HBS and the complexity of HBS patches on the fire landscape. As maximum air temperatures for the month of ignition approached 40^o C, the percentage of HBS to total area burned frequently exceeded 20%. The percentage of HBS to total area burned also exceed 20% when the precipitation total recorded during the previous 12 months was less than 25% of the annual average precipitation. Landscape analysis results showed that, as the total area burned in fires on the Central California coast grows, the edge lengths and areas of HBS patches also grows at a rapid rate. At the same time, the perimeter-to-area ratio of HBS patches decreases gradually and the HBS patches become more aggregated as total burned area grows.     

Prevalence, patterns, and effects of shell damage on Geukensia demissa in South Carolina estuarine habitats

Shelled molluscs frequently exhibit a record of damage on exterior surfaces that can evidence past predation attempts and may affect survival and growth. In South Carolina populations of the ribbed marsh mussel, Geukensia demissa, >90% of the individuals and up to 60% of the total shell area are damaged. A trend toward greater amounts of damage occurred on mid-marsh compared to oyster reef mussels from the barrier beach side of inlets. Shell damage effects on survivorship and shell and tissue growth were assessed seasonally during multi- and single-season field experiments. Mussels from a common mid-marsh site were divided into size classes (~50 or 70 mm), treated to create two damage levels (undamaged and damaged), and replaced within mid-marsh exclusion cages to minimize additional shell damage. In both multi- and single-season experiments increased shell damage resulted in significantly greater mortality. Linear shell growth was unaffected by increased damage, but 50 mm mussels grew twice as fast. Shell mass increased 16–50% in the multi-season and single-season winter period, but decreased 7–12% during the single-season summer period. Tissue mass significantly decreased 31–43% in 50 mm damaged mussels, but increased by 33% for 70 mm mussels in both multi-season and the single-season winter period experiments. Shell damage did reduce tissue mass 43% in 70 mm single-season summer mussels. Experimental results indicate shell damage from a simulated increase in predation can affect negatively both survival and growth of marsh mussels. Seasonal timing of shell damage and initial mussel size also influenced the effects of sublethal predation on shell and tissue growth. The previously unrecognized importance of sublethal predation and the resultant significant negative effects of shell damage on survival and growth will affect the distribution and population dynamics of G. demissa in coastal marshes and will influence the overall contribution of ribbed mussels to estuarine ecosystems.     

Carry-over effects of male extra-pair copulation opportunity on biparental effort in zebra finches

Whether parental effort can be negotiated between partners over ecological time and adjusted across different contexts is not well understood. We manipulated male extra-pair copulation (EPC) opportunity in captive zebra finches, Taeniopygia guttata, to test whether males adjust incubation effort to the mating context and to examine how females respond to their partner’s effort. Birds without previous breeding experience were paired randomly and bred with the same partner twice. In the first breeding attempt, half the males received EPC opportunities with ‘extra-pair females’ during incubation, while the other half did not. Males that received EPC opportunities in the first breeding attempt did not in the second breeding attempt and vice versa. We recorded incubation effort on days when EPC opportunities were not presented. In their first breeding attempt, males with EPC opportunities incubated less than those without. Females compensated fully for the deficit in male care so that a pair’s combined incubation effort was unchanged. In the second attempt, when a male’s opportunity for EPCs was switched, individuals showed the same level of incubation effort that they had previously, irrespective of the current availability of extra-pair females. This suggests that division of effort was negotiated in the first breeding attempt and maintained without significant adjustments in the second attempt. The effects of male EPC opportunity in the first breeding attempt on subsequent incubation effort suggests that individual parental decisions can be shaped by previous experience and this may partly explain conflicting results in studies where individuals’ histories were not known.     

Correlates of male mating success in great bustard leks: the effects of age, weight, and display effort

We examined how mating success varied in relation to age, weight, body size, and display behavior among great bustard Otis tarda males. The estimated mating success was strongly skewed, with 45% of adult males being involved in copulation attempts and only 9.7% actually seen copulating successfully. Unlike most birds, body size continued increasing in great bustards several years after reaching sexual maturity. Age, weight, and display effort were all significant and independent predictors of male mating success. The higher display effort involved performing longer full-display bouts. Older males could detach from the male flock earlier in the season as well as on each day and spend longer seasonal and daily periods displaying as solitary birds, which contributed to increase their mating success. In contrast, males weighing more did not invest more in display, which suggests that they could be recognized as dominants by other males and selected by females through assessment of their plumage sexual traits. In contrast to most other bird species, the system described for great bustards resembles that found in some lek-mating ungulates, where social rank is a complex trait determined by both age and mass, and as in these mammals, it suggests that sexual selection continues to favor a high male weight in this extremely sexually dimorphic species.     

Differential effects of energy and mass influx on the landscape synchrony of lake ecosystems

Interannual variation of 45 annually resolved time series of environmental, limnological, and biotic parameters was quantified (1994-2009) in six lakes within 52,000 km2 to test the hypothesis that influx of energy (E; as irradiance, heat, wind) varies synchronously among sites and induces temporal coherence in lakes and their food webs, whereas influx of mass (m; as water, solutes, particles) reduces synchrony because local catchments uniquely modify hydrologic inputs. Overall, 82% of parameters exhibited significant (P < 0.05) synchrony (S) estimated as mean pair-wise correlation of Z-transformed time series. Influx of E as atmospheric heat and irradiance was both more highly synchronous and less temporally variable (months-to-decades) than influx of m as summer precipitation, snow, or river discharge. Similarly, S of limnological parameters varied from 0.08 to 0.85, with variables known to be regulated by E influx (ice melt, gas solubility) up to twofold more coherent than those regulated by m inputs (organic solutes). Pairs of variables linked by simple direct mechanisms exhibited similar S values (air temperature and ice melt, nutrients and algae), whereas the coherence of other parameters (water temperature, mixing) was intermediate to that of multiple regulatory agents. Overall, aggregate measures of plankton density varied more coherently among lakes than did constituent taxa. These findings suggest that environmental variability is transmitted to most levels of aquatic ecosystems, but that the precise effects depend on whether E or m fluxes predominate, the coherence of each forcing mechanism, and the strength of linkages between exogenous forcing and lake response.     

Seasonal density dependence, timing of mortality, and sustainable harvesting

Birth-pulse populations are often characterized with discrete-time models, that use a single function to relate the post-breeding population size to the post-breeding size of the previous year. Recently, models of seasonal density dependence have been constructed that emphasize interactions during shorter time periods also. Here, we study two very simple forms of density-dependent mortality, that lead to Ricker and Beverton-Holt type population dynamics when viewed over the whole year. We explore the consequences of harvest timing to equilibrium population sizes under such density dependence. Whether or not individual mortality compensates for the harvested quota, the timing of harvesting has a strong impact on the sustainability of a harvesting quota. Further, we show that careless discretization of a continuous mortality scheme may seriously underestimate the reduction in population size caused by hunting and overestimate the sustainable yield. We also introduce the concept of the demographic value of an individual, which reflects the expected contribution to population size over time in the presence of density dependence. Finally, we discuss the possibility of calculating demographic values as means of optimizing harvest strategies. Here, a Pareto optimal harvest strategy will minimize the loss of demographic value from the population for a given yield.     

Plant diversity partitioning in Mediterranean croplands: effects of farming intensity, field edge, and landscape context

Farmland biodiversity is affected by factors acting at various spatial scales. However, most studies to date have focused on the field or farm scales that only account for local (alpha) diversity, and these may underestimate the contribution of other diversity components (beta diversity) to total (gamma) farmland diversity. In this work, we aimed to identify the most suitable management options and the scale at which they should be implemented to maximize benefits for diversity. We used a multi-scale additive partitioning approach, with data on plant diversity from 640 plots in 32 cereal crop fields from three agricultural regions of central Spain that differed in landscape configuration. We analyzed the relative contribution to overall plant diversity of different diversity components at various spatial scales and how these diversity components responded to a set of local (application of agri-environment schemes [AES] and position within the field) and landscape (field size and landscape connectivity and composition) factors. Differences in species composition among regions and then among fields within regions contributed most to overall plant diversity. Positive edge effects were found on all diversity components at both the field- and regional scales, whereas application of AES benefited all diversity components only at the field scale. Landscape factors had strong influences on plant diversity, especially length of seminatural boundaries, which increased species richness at both the field and the regional scales. In addition, positive effects of percentage of nonproductive land-uses in the landscape were found on all diversity components at the regional scale. Results showed that components that contributed most to overall plant diversity were not benefited by current AES. We conclude that agri-environmental policies should incorporate and prioritize measures aimed at the maintenance of seminatural boundaries and patches of nonproductive habitats within agricultural landscapes, through landscape planning, cross-compliance, or high nature-value farmland programs. These options will help to conserve overall plant diversity at regional scales, as well as the spillover of plant species from such seminatural elements into crops, especially in Mediterranean areas that still harbor extensive farming and relatively complex landscapes.     

Within-brood patterns of paternity and paternal behavior in red-winged blackbirds

We video-taped male and female red-winged blackbirds (Agelaius phoeniceus) feeding individual chicks in order to test the hypothesis that food might be differently allocated to within-pair offspring and extra-pair young. We found no evidence that paternity influenced the allocation of food by either males or females. Both males and females fed male offspring significantly more, but there was no tendency for paternity to be skewed by gender. Females fed older offspring significantly more, whereas males did not; extra-pair fertilizations, however, were not associated with lay or hatch order of the chicks. Given that males do not appear to discriminate within-pair from extra-pair offspring directly, these results are consistent with current theory on the effect of paternity on paternal behavior. We discuss briefly some of the possible reasons why discrimination might be lacking in red-winged blackbirds and in other species in which the possibility of discrimination of paternity and allocation of paternal behavior has been studied.     

Inter- and intraspecific patterns of reproductive effort in four species of cone shells (Conus spp.)

Patterns of reproductive effort were examined both within and between 4 species of the tropical marine gastropod genus Conus from Hawaii in 1979. Caloric content was measured for the somatic tissues of female cone shells and for their egg capsule masses. These data were used to calculate (1) current reproductive effort, which was the ratio of annual energy produced as spawn to the sum of the annual energy produced as spawn plus the annual energy produced as somatic tissue; and (2) weight-specific reproductive effort, which was the ratio of annual energy produced as spawn to the total energy content of the female body just before spawning. For C. pennaceus, during the approximately 10 yr of life of a female, current reproductive effort rose from about 0.35 at 2 yr to about 0.85 at 10 yr. In contrast, weight-specific reproductive effort remained relatively constant throughout life for C. pennaceus (about 0.2 to 0.5), C. abbreviatus (about 0.2 to 0.3), C. flavidus (about 0.2 to 0.3) and C. quercinus (about 0.1 to 0.2). It is suggested that weight-specific reproductive effort may be useful as an index of the selective importance of the survival cost of present reproduction, while current reproductive effort more closely reflects the selective importance of energy allocation tradeoffs. Data on the 4 species of Conus examined support the hypothesis that reproductive effort should be positively correlated with extrinsic adult mortality and negatively correlated with the variability of juvenile survivorship. The data also suggest a possible relationship between reproductive effort and the cost of parental care.     

Settlement patterns and presettlement behavior of the naked goby,Gobiosoma bosci,a temperate oyster reef fish

Dispersion, distribution, development and feeding incidence of larvae of the naked goby,Gobiosoma bosci (Lacepéde), were examined for linkages between larval behavior while near the reef surface and later patterns of settlement and recruitment. Field sampling and experiments were conducted during the summers of 1988 and 1989 in the Flag Pond oyster reef along the western shore of the Chesapeake Bay near Camp Conoy, Maryland, USA. Results indicated that prior to settlement most demersal larvae aggregate in shoals and exhibit distinct microhabitat preferences on the reef. In a field experiment, larvae settled both during the day and at night. Dispersion at settlement was aggregated, suggesting that demersal shoaling influences settlement patterns in this species. The distribution of demersal larvae also indicated that larval swimming behavior is sufficiently strong to permit active control of position on reefs. Large demersal larvae settled rapidly when brought to the laboratory, but small larvae in demersal shoals appeared to require additional growth and morphological development prior to settlement. Development of the pelvic fins, used by juveniles and adults for perching on the substrate, may be a good indicator of competence to settle in this species. The adaptive significance of demersal shoaling by small larvae of the naked goby, and the fate of these larvae, remains perplexing because the low feeding rates found for larvae shoaling near the reef surface should slow or prevent the growth and development required prior to settlement. Observations made by other authors indicate that demersal shoaling and the use of water directly overlying reefs may be common behaviors of temperate and tropical reef fishes.     

Associations of forest bird species richness with housing and landscape patterns across the USA.

In the United States, housing density has substantially increased in and adjacent to forests. Our goal in this study was to identify how housing density and human populations are associated with avian diversity. We compared these associations to those between landscape pattern and avian diversity, and we examined how these associations vary across the conterminous forested United States. Using data from the North American Breeding Bird Survey, the U.S. Census, and the National Land Cover Database, we focused on forest and woodland bird communities and conducted our analysis at multiple levels of model specificity, first using a coarse-thematic resolution (basic models), then using a larger number of fine-thematic resolution variables (refined models). We found that housing development was associated with forest bird species richness in all forested ecoregions of the conterminous United States. However, there were important differences among ecoregions. In the basic models, housing density accounted for < 5% of variance in avian species richness. In refined models, 85% of models included housing density and/or residential land cover as significant variables. The strongest guild response was demonstrated in the Adirondack-New England ecoregion, where 29% of variation in richness of the permanent resident guild was associated with housing density. Model improvements due to regional stratification were most pronounced for cavity nesters and short-distance migrants, suggesting that these guilds may be especially sensitive to regional processes. The varying patterns of association between avian richness and attributes associated with landscape structure suggested that landscape context was an important mediating factor affecting how biodiversity responds to landscape changes. Our analysis suggested that simple, broadly applicable, land use recommendations cannot be derived from our results. Rather, anticipating future avian response to land use intensification (or reversion to native vegetation) has to be conditioned on the current landscape context and the species group of interest. Our results show that housing density and residential land cover were significant predictors of forest bird species richness, and their prediction strengths are likely to increase as development continues.     

Serial patterns of biodiversity change in corals across shallow reef flats in Ko Phuket,Thailand, due to the effects of local (sedimentation) and regional (climatic) perturbations

A 17-year monitoring programme of reef flats at Ko Phuket, Thailand afforded an opportunity to evaluate both univariate and multivariate measures of environmental stress in an assessment of change on coral reef ecosystems. The sites at Ko Phuket suffered the effects of dredging in 1986-1987 and then anomalously low sea levels in 1997-1998 as a result of climate-related events in the Indian Ocean. Univariate measures of species diversity and taxonomic distinctness ((*) increased across the reef flat, reflecting the greater effects of physical stresses on the inner reef flats (compared with outer reef flats) at all sites, with more congeneric species present on the inner reef and more confamilial species on the outer reef. Multivariate measures showed a clear breakdown in seriation patterns at all sites during the dredging in 1987 and in 1998, as a result of earlier negative sea-level anomalies. Recovery from environmental disturbances was obvious within 12 months in each case. The domination of the reefs by massive coral species, which are physiologically adapted to intertidal living and which display partial rather than total colony mortality, may be a factor contributing to the apparent resilience of the reef flats together with continued recruitment and survival of juvenile corals during adverse environmental conditions. Elevated sea temperatures and extensive bleaching of corals in 1991, 1995, and 1998 had no effect upon coral community measures, with many corals recovering their zooxanthellae numbers within 3-5 months of the bleaching events.