Linearity in the aggregate effects of multiple predators in a food web

Theory in community ecology often assumes that predator species have similar indirect effects and thus can be treated mathematically as a single functional unit (e.g. guild or trophic level). This assumption is questionable biologically because predator species typically differ in their effects, creating the potential for nonlinearities when they coexist. We evaluated the nature of indirect effects caused by three species of hunting spider predators, singly and in multiple species combinations, on grass and herb plants in experimental old-field food webs. Despite the potential for nonlinearity, indirect effects in different multiple predator combinations consistently did not differ significantly from the respective means of the single species effects. Thus, for this experimental system, the whole was simply the average of the parts. Consequently, models which abstract predator species as single trophic levels would successfully predict indirect effects in this system regardless of the composition of the predator fauna.     

Cooccurrence of prey species alters the impact of predators on prey performance through multiple mechanisms

When prey are differentially affected by intra and interspecific competition, the cooccurrence of multiple prey species alters the per capita availability of food for a particular prey species which could alter how prey respond to the threat of predation, and hence the overall‐effect of predators. We conducted an experiment to examine the extent to which the nonconsumptive and overall effect of predatory water bugs on snail and tadpole traits (performance and morphology) depended on whether tadpoles and snails cooccurred. Tadpoles and snails differed in their relative susceptibility to intraspecific and interspecific competition, and predators affected both prey species via consumptive and nonconsumptive mechanisms. Furthermore, the overall effect of predators often depended on whether another prey species was present. The reasoning for why the overall effect of predators depended on whether prey species cooccurred, however, differed for each of the response variables. Predators affected snail body growth via nonconsumptive mechanisms, but the change in the overall effect of predators on snail body growth was attributable to how snails responded to competition in the absence of predators, rather than a change in how snails responded to the threat of predation. Predators did not affect tadpole body growth via nonconsumptive mechanisms, but the greater vulnerability of competitively superior prey (snails) to predators increased the strength of consumptive mechanisms (and hence the overall effect) through which predators affected tadpole growth. Predators affected tadpole morphology via nonconsumptive mechanisms, but the greater propensity for predators to kill competitively superior prey (snails) enhanced the ability of tadpoles to alter their morphology in response to the threat of predation by creating an environment where tadpoles had a higher per capita supply of food available to invest in the development of morphological defenses. Our work indicates that the mechanisms through which predators affect prey depends on the other members of the community.     

Habitat selection within species ofDrosophila: a review of experimental findings

Summary Habitat selection whereby individuals within a population have different microhabitat preferences is possibly important for genetic diversity and stability. It has received a good deal of attention, especially forDrosophila populations. The literature is summarized and an attempt is made to reconcile what appear to be contradictory findings. Some generalizations are drawn: (1) individualDrosophila choose habitats in consistent and characteristic ways; (2) some microhabitat differences are more important to flies than others; (3) genetic factors influence habitat preferences; (4) experiential factors influence habitat preferences; (5) differences in processing ability moderate the influence of experience on habitat choice. It is argued that the traditional analysis of variance models are frequently not very helpful for visualizing the process, and a computational model is suggested instead.     

Dying from the lesser of three evils: facilitation and non‐consumptive effects emerge in a model with multiple predators

Prey modify their behaviour to avoid predation, but dilemmas arise when predators vary in hunting style. Behaviours that successfully evade one predator sometimes facilitate exposure to another predator, forcing the prey to choose the lesser of two evils. In such cases, we need to quantify behavioural strategies in a mix of predators. We model optimal behaviour of Atlantic cod Gadus morhua larvae in a water column, and find the minimal vulnerability from three common predator groups with different hunting modes; 1) ambush predators that sit‐and‐wait for approaching fish larvae; 2) cruising invertebrates that eat larvae in their path; and 3) fish which are visually hunting predators. We use a state‐dependent model to find optimal behaviours (vertical position and swimming speed over a diel light cycle) under any given exposure to the three distinct modes of predation. We then vary abundance of each predator and quantify direct and indirect effects of predation. The nature and strength of direct and indirect effects varied with predator type and abundance. Larvae escaped about half the mortality from fish by swimming deeper to avoid light, but their activity level and cumulative predation from ambush predators increased. When ambush invertebrates dominated, it was optimal to be less active but in more lit habitats, and predation from fish increased. Against cruising predators, there was no remedy. In all cases, the shift in behaviour allowed growth to remain almost the same, while total predation were cut by one third. In early life stages with high and size‐dependent mortality rates, growth rate can be a poor measure of the importance of behavioural strategies.     

Habitat selection,interspecific interactions and landscape composition

Summary I argue here that, from the perspective of any individual, most landscapes are composed of only three basic types of habitats. These are: (1) source habitat in which reproduction exceeds mortality and the expected per capita growth rate is greater than one; (2) sink habitat, in which limited, reproduction is possible but will not on average, compensate for mortality and the per capita rate of growth is between zero and one; and (3) unusable habitat, which comprises the matrix of all habitats that are never exploited by the species in question, and in which patches of source and sink habitats are embedded. Unlike earlier source-sink models, this model explicitly considers the effects that substituting one type of habitat for another has on the equilibrium size of a population and the interactions between species which can use both source and sink habitats. The model demonstrates that the equilibrium size of a species' population can sometimes be increased by substituting unusable habitat for sink habitat. Thus, even though the average patch quality in the landscape may be decreased, the overall quality of the landscape can increase. For two species with distinct habitat preferences, interactions between species can vary qualitatively as well as quantitatively as a function of the relative abundances of each of the habitat types. The model also shows that the interactions between species are particularly sensitive to the relative costs of moving between patches and sampling patches to determine their quality. Recent fragmentation of natural landscapes may increase the cost of searching for usable (source or sink) patches. Under some conditions, the interspecific interactions may be substantially more negative (competitive) than the interactions that evolved in the original natural landscape, further reducing population sizes and increasing the likelihood of competitive exclusion in fragmented modern landscapes.     

Foraging distribution overlap and marine reserve usage amongst sub-Antarctic predators inferred from a multi-species satellite tagging experiment

Satellite telemetry data was used to predict at sea spatial usage of five top order and meso-predators; Antarctic fur seals (Arctocephalus gazella), macaroni penguins (Eudyptes chrysolophus), king penguins (Aptenodytes patagonicus), black browed albatross (Diomedea melanophrys), and light mantled albatross (Phoebetria palpebrata). All were tagged at Heard Island in the Southern Ocean over a single summer season collecting over 5000 tracking days for 178 individuals. We aimed to predict areas of likely high foraging value from tracking environmental data and also to quantify overlap in foraging range between species. Hidden Markov models were used to differentiate between bouts of Area Restricted Search (ARS) assumed to be associated with areas of higher foraging value, and transit behaviours. Oceanographic and distance metrics associated with ARS activity were then used to calculate a habitat electivity index. A combined bootstrap/Monte Carlo scheme was employed to propagate uncertainty from the Hidden Markov models into the habitat prediction scheme. Distinct differences were predicted in the spatial distribution of foraging locations in different species, reflecting different dispersive abilities and foraging strategy. The extent of usage and foraging distribution was largely contained within Australian the Economic Exclusion Zone (EEZ). In comparison, the smaller Australian Commonwealth Marine Protected Areas (MPAs) contained <20% of the predicted foraging distributions.     

动物生境选择研究中的时空尺度

尺度研究已成为生态学上的一个重要概念和研究热点,但是在动物生境选择的研究中尚未引起足够的重视。动物的生境选择包括多层次的判别和一系列等级序位,在各个尺度和水平上具有不同的特征和机制,受到时空尺度的严格限定。繁殖期的时间限制、社群压力、环境变化、动物生理需求的变化决定了动物生境选择的时间制约性;而生境资源的斑块化分布、功能生境之间的相互作用决定了动物生境选择的空间制约性。研究者对时间和空间尺度的选取与应用会直接影响到生境选择研究结果的科学性和实效性。本文从动物生境选择的时空制约性出发,分析了生境选择研究中时间尺度的重要性,叙述了国内外生境选择研究中常见的研究尺度,强调了多尺度研究和长期生态研究的必要性,尺度的选择应该成为生境选择研究的起点和基础。     

Habitat selection of juvenile banana prawns, Penaeus merguiensis de Man: Testing the roles of habitat structure, predators, light phase and prawn size

The effects of fish predators, light phase, habitat structure and prawn size on the habitat preferences of juvenile Penaeus (Fenneropenaeus) merguiensis de Man were examined with laboratory experiments. The behaviour of juvenile P. merguiensis within habitats of different structural complexity was also examined. Experiments were carried out in a tank (1.8 m diameter) divided into four habitats representing: bare substratum, leaf litter (little vertical structure), mangrove pneumatophores (regular vertical structure) and mangrove woody debris (heterogeneous vertical structure). The location of 10 prawns was monitored over 270 min (135 min light and 135 min dark), with different prawns five times for each combination of prawn size class, and predator (no predator; Arius graeffei Kner and Steindachner and Lates calcarifer Bloch).In the absence of predators and during the light phase, when observations on prawn behaviour were made, swimming was the most common behaviour (of seven mutually exclusive behavioural categories) with few differences in behaviour between sizes. All size classes of juvenile P. merguiensis selected vertical structure (mangrove debris and pneumatophores) over low vertical structure (leaf litter and bare substratum), in both light and dark conditions and in the presence or absence of predators. When L. calcarifer was present, the selection by prawns of the mangrove-debris habitat increased significantly. This was attributed to an increase in predation risk in the other habitats. L. calcarifer rarely pursued prey amongst the mangrove-debris structure, compared to habitats with less heterogeneous vertical structure (pneumatophores, leaf litter and bare substratum).     

Habitat selection and subsequent reproductive success in the beaugregory damselfish

Synopsis Male beaugregory damselfish,Stegastes leucostictus, were provided with three types of artificial breeding structures to determine if they change habitats based on past or future reproductive payoffs. All three site types quickly lured males away from their natural sites. In comparison to those living on natural sites, those using artificial sites were less likely to move to different areas and had a higher reproductive success. When given no choice, male reproductive success was correlated to structural type. A second experiment provided males with an additional structure after using the intermediate quality type for 2 months. Males would often initially use both sites but would eventually shift their spawning activity to the new site if it was of the same quality or better than the old one. However, males would not move if the new site was of inferior quality. When given a new site identical to the initial one, approximately half of the males shifted to the new site. There was no evidence that reproductive performance influenced a male's decision to use a new site.     

Foraging in a landscape mosaic: selection for energy and minerals in free-ranging cattle

Several studies have indicated the potential importance of nutrients, other than energy, in determining foraging decisions. A model was developed to test this idea, on the assumption of an intake maximization for different nutrients (energy, sodium and phosphorus). The model predictions were tested using field data from cattle grazing in a landscape mosaic of Pleistocene cover-sand and riverine grassland. Observations on foraging behaviour, food intake and diet composition were collected in thirteen 4-day-periods over 2 years. Habitat selection was determined by comparing the proportion of grazing time in different vegetation units with the available area proportion of the units. Two levels of habitat selection were examined: a micro-level (fine-scale, where vegetation units were considered separately) and a macro-level (coarse-scale, where vegetation units were combined to give selection at the landscape level). At the micro-level of habitat selection, no selection was apparent between the vegetation units of the riverine landscape, but the Deschampsia flexuosa unit was significantly selected for in the cover-sand landscape. At the landscape (macro-) level, the animals preferred the riverine landscape. The model revealed poor predictions of habitat occupancy on a micro-level. A much better prediction was obtained when vegetation units were combined at a macro-level. The D. flexuosa unit provided a higher energy intake, whereas the intake of sodium was higher in riverine grassland. Phosphorus proved relatively significant in determining habitat occupancy. Based on energy maximization alone, the model was a very poor predictor of habitat occupancy. It is argued that selection occurred mainly at the macro-level. The incorporation of different nutrient constraints in foraging models can then prove fruitful when seeking explanations of habitat occupancy. At the micro-level, difficulties for the animal in assessing nutrient availability may result in a less selective foraging pattern. However, the costs of increased selectivity may be greater than the benefits.     

Habitat selection behaviour links local and regional scales in aquatic systems

The role of habitat selection behaviour in the assembly of natural communities is an increasingly important theme in ecology. At the same time, ecologists and conservation biologists are keenly interested in scale and how processes at scales from local to regional interact to determine species distributions and patterns of biodiversity. How important is habitat selection in generating observed patterns of distribution and diversity at multiple spatial scales? In theory, habitat selection in response to interacting species can generate both positive and negative covariances among species distributions and create the potential to link processes of community assembly across multiple scales. Here I demonstrate that habitat selection by treefrogs in response to the distribution of fish predators functions at both the regional scale among localities and the local scale among patches within localities, implicating habitat selection as a critical link between local communities and the regional dynamics of metacommunities in complex landscapes.     

四川短尾鼩的生境选择及对人为干扰的响应

2014年4-9月,通过铗夜法对12种不同生境内的四川短尾鼩种群数量进行了调查,依据捕获率的大小及生境的干扰程度分析四川短尾鼩的生境选择倾向。利用人工捕捉模拟种群干扰,分别在干扰1个月、2个月、3个月、4个月时间之后,监测荒草地、废弃梨树林、灌丛和绿化林4种代表性生境内四川短尾鼩的种群数量、年龄结构、性比的变化,进而分析四川短尾鼩对人为干扰的生态响应。结果显示：（1）四川短尾鼩在12种典型生境中,按捕获率的大小依次分为4个等级：绿化林＞废弃梨树林、景观林、荒草地、桉树林、蔬菜地、灌丛＞人居村落、废弃葡萄园＞竹林,葡萄园,油菜地;（2）四川短尾鼩的生境选择倾向为：异质性适中、适度干扰、竞争较低的生境＞异质性较低、干扰较高的生境（宜觅食、竞争弱）或异质性较高、干扰较低的生境（宜栖身、竞争弱）＞高竞争的生境＞异质性低或强烈干扰或二者兼有的生境。（3）干扰之初,所有生境中四川短尾鼩种群密度明显降低,干扰2个月之后其种群数量均可恢复到正常状态;干扰后性比平均值增加3.52%,怀孕率变化不显著;全部生境中,成体组比例平均降低1.71%,老年组减少7.53%;幼年组的比例平均增长2.33%,亚成体增加6.91%。幼体和亚成体比例的增加,使种群数量在一定时间内保持稳定增长。     

Habitat use by Oncifelis guigna and Pseudalopex culpaeus in a fragmented forest landscape in central Chile

Native forests in Chile have been replaced by exotic plantations, generating a mosaic of forest fragments immersed in a matrix of plantations and farms. This landscape change could affect habitat availability for native carnivores such as Oncifelis guigna (Felidae) and Pseudalopex culpaeus (Canidae). We determine the potential impact of native forest fragmentation on these species, assessinghabitat use by these carnivores. The study was carried out in continuous forest (150 ha), native forest fragments (<10 ha) and surrounding pine plantations in central Chile (35°59 S, 72°41 W). We placed scent-stations (SSs) in three types of habitat from January to August 2000, setting a total of 620 SSs to determine habitat selection. We characterized structure and composition of 100 m² of vegetation aroundevery SS, and measured its distance to nearest native forest patch, road and bottom creek. O. guigna preferred habitats with dense bush cover, far from roads and close to large patches of native forest, being almost exclusively restricted to this type of habitat. On the other hand, P. culpaeus preferred openhabitats, close to roads, relying extensively on pine plantations but scarcely using forest and forest fragments. Our results suggest that native forest fragmentation, considering habitat use only, would have a negative effect on O. guigna populations. Thus, forest size reduction and isolation may reduce O.guigna survival in a pine dominated landscape. In contrast, P. culpaeus, a generalist species that would use habitat depending on its abundance, may adapt to changes in this landscape.     

Nonconsumptive effects in a multiple predator system reduce the foraging efficiency of a keystone predator

Many studies have demonstrated that the nonconsumptive effect (NCE) of predators on prey traits can alter prey demographics in ways that are just as strong as the consumptive effect (CE) of predators. Less well studied, however, is how the CE and NCE of multiple predator species can interact to influence the combined effect of multiple predators on prey mortality. We examined the extent to which the NCE of one predator altered the CE of another predator on a shared prey and evaluated whether we can better predict the combined impact of multiple predators on prey when accounting for this influence. We conducted a set of experiments with larval dragonflies, adult newts (a known keystone predator), and their tadpole prey. We quantified the CE and NCE of each predator, the extent to which NCEs from one predator alters the CE of the second predator, and the combined effect of both predators on prey mortality. We then compared the combined effect of both predators on prey mortality to four predictive models. Dragonflies caused more tadpoles to hide under leaf litter (a NCE), where newts spend less time foraging, which reduced the foraging success (CE) of newts. Newts altered tadpole behavior but not in a way that altered the foraging success of dragonflies. Our study suggests that we can better predict the combined effect of multiple predators on prey when we incorporate the influence of interactions between the CE and NCE of multiple predators into a predictive model. In our case, the threat of predation to prey by one predator reduced the foraging efficiency of a keystone predator. Consequently, the ability of a predator to fill a keystone role could be compromised by the presence of other predators.     

Guild‐level responses by mammalian predators to afforestation and subsequent restoration in a formerly treeless peatland landscape

Afforestation of formerly open landscapes can transform mammalian predator communities, potentially impacting prey species like ground‐nesting birds. In Scotland's Flow Country, a globally important peatland containing many forestry plantations, earlier studies found reduced densities of breeding waders on open bogs, when forestry plantations were present within 700 m. One plausible explanation for this pattern is mammalian predation. We tested whether mammalian predator indices, based on scats (feces), differed between (1) open bog, forestry plantations, and former plantations being restored as bog (“restoration” habitats); (2) restoration habitats of different ages; and (3) open bogs with differing amounts of nearby forestry. We measured summer scat density and size over 14 years in 26 transects 0.6–4.5 km in length, collecting data during 93, 96, and 79 transect‐years in bog, forestry, and restoration habitats respectively. In forestry, scat density increased eightfold, reaching values ~6 times higher than those of bogs. On open bogs with over 10% forestry within 700 m, scat densities were 2.9 times higher than on open bogs with less forestry nearby. Results support the hypothesis that mammalian predators might be responsible for the low densities of breeding waders close to forests, on adjacent open bogs. In restoration habitats, scat densities rose 6–10 years after felling but fell to levels similar to open bogs in older restoration habitats, supporting restoration management as a means of reducing mammalian predator activity/abundance. We urge caution around decisions to establish forestry plantations in open landscapes of high biodiversity importance.     

Importance of visual cues of conspecifics and predators during the habitat selection of coral reef fish larvae

The study investigated visual recognition of conspecifics and predators by settlement-stage coral reef fish larvae in a set of three experiments using a dual-choice aquarium (Moorea Island). Experiments 1 and 2 were conducted under artificial light conditions. Experiment 3 was conducted under natural light during new and full moon nights. In experiment 1, five out of six species preferred conspecifics rather than heterospecifics (Acanthurus triostegus, Chromis viridis, Ostorhinchus angustatus, Stegastes fasciolatus, Valenciaenna strigata). In experiment 2, three out of six species were repulsed by predators (Mulloidichtys flavolineatus, O. angustatus, V. strigata). In experiment 3 (conducted on one species), A. triostegus was attracted to conspecifics during bright nights, but did not show such behavior during dark nights. Our study raises the question of trade-off for fish larvae to settle during the night with high light intensities to favor the visual recognition of conspecifics and predators, or during darker nights to reduce reef predation.     

四川山鹧鸪的分布及生境选择

研究了四川山鹧鸪(Arborophila rufipectus)的分布区域和栖息地的生境,认为四川山鹧鸪的分布范围比已知的要大,但其实际分布区呈明显的岛屿状,显示出生境的破碎化;指出四川山鹧鸪的适宜生境是原始的常绿阔叶林、针阔混交林和具有较大常绿落叶阔叶乔木树种盖度的多年生次生林,亦可选择部分人工林,而对地表灌丛密度大的次生幼林以及人工幼林生境不喜好。由于天然林的禁伐和生态林的管护,四川山鹧鸪的栖息地趋于稳定并有所扩大。但四川山鹧鸪仍然处于濒危状态。     

西藏慈巴沟自然保护区羚牛栖息地选择

2001 年12 月至2002 年8 月,笔者在西藏慈巴沟自然保护区对羚牛栖息地选择性及其季节变化进行了调查研究。调查中,沿慈巴沟河谷随机选择起始点,系统地设置了187个10 m×10 m 的固定样方,记录了样方所在位置的植被类型、海拔、坡度、坡向、距水源距离、乔木密度、灌木密度、箭竹密度、郁闭度和隐蔽度等,并分别在冬、春、夏三季检查样方是否被羚牛利用。本文通过计算Vanderloeg 选择系数和Scavia 选择指数来研究羚牛对植被的选择性,结果发现羚牛冬季对杜鹃针叶林、箭竹针叶林和针阔混交林具有正选择性,春季则对箭竹针叶林和针阔混交林、夏季对高山灌丛草甸和杜鹃针叶林具有正选择性。对9 个数量化因子进行分组处理,利用Vanderloeg 选择系数和Scavia 选择指数,计算了这些因子在各个季节不同的选择性,进一步分析了羚牛对栖息地的选择性。对9 个数量化因子的判别分析表明,冬、夏及春、夏之间有极显著差异,海拔对判别函数的贡献最大,其它因子如箭竹密度、隐蔽度、郁闭度和乔木密度等贡献也较大,说明这些因子的季节分离明显。而羚牛的冬、春之间判别分析不显著,季节分离的程度较小。在结合其它分布区羚牛栖息地研究的基础上,本文探讨了食物和盐源在羚牛栖息地选择季节变化中的作用。     

Habitat selection rules in breeding mallards (Anas platyrhynchos): a test of two competing hypotheses

Ideal preemption and conspecific attraction are alternative hypotheses of the habitat selection rules used by individuals. According to the former an occupied site is assumed to be preempted and therefore not available for later arriving individuals, whereas according to the latter individuals are assumed to be attracted by conspecifics to occupied sites, rather than avoiding them. We studied these competing hypotheses in breeding mallards (Anas platyrhynchos) by a cross-over experiment in 2 years, introducing birds onto lakes before migratory wild mallards arrived. If mallards use the ideal preemptive rule, breeding density of wild mallards in experimental lakes should be lower and they should be occupied less frequently than control lakes, but if mallards use the conspecific attraction rule the reverse should be true. Our results allowed us to reject the ideal preemptive rule whereas the conspecific attraction rule was to some extent supported. We discuss these findings in relation to population limitation. The results suggest that the local breeding population studied is not limited by spacing behaviour related to habitat selection. Received: 14 July 1997 / Accepted: 17 November 1997     

Measures of giant panda habitat selection across multiple spatial scales for species conservation

Examining ecological processes across spatial scales is crucial as animals select and use resources at different scales. We carried out field surveys in September 2005, March–September 2006, and April 2007, and used ecological niche factor analysis to determine habitat preferences for the giant panda (Ailuropoda melanoleuca) across 4 spatial scales: daily movement, core range, home range, and seasonal elevational migration. We found that giant pandas prefer conifer forest and mixed forest at higher than average elevation (2,157 m) of study area in the 4 scale models. However, we also observed significant scale differences in habitat selection. The strength of habitat preference increased with scale for the 2 disturbed forests (sparse forest and fragmented forest), and decreased with scale for 0–30° gentle slope and south- and north-facing aspect. Furthermore, habitat suitability patterns were scale-dependent. These findings highlight the need to determine species–environment associations across multiple scales for habitat management and species conservation. © 2012 The Wildlife Society.