Pollinator functional response and plant population dynamics: Pollinators as a limiting resource

Summary During recent years, much work has focused on which factors limit the reproductive success in plants. Several studies show a strong influence of either resource limitation, pollen limitation or a combined effect of both. The theoretical arguments for resource limitation are abundant, but there has been very little work done concerning the effect of pollinator availability. In this paper we construct a model to study how the reproductive success in plants is influenced by the foraging behaviour of the pollinators. The pollinator population is assumed to have a constant population density. A functional response function for the pollinators is derived. It is similar to a Holling type II functional response. It is shown that, since the pollinators are regulated by factors not included in the model and their capability to pollinate is limited by the functional response, this is sufficient for regulating the plant population. There also exists a threshold condition for the persistence of the plant population that depended on the search rate of the pollinators and the demographic parameters of the plant population. If this threshold condition is not satisfied the plant population cannot persist and will become extinct. If the condition is satisfied the plant population grows until it is limited at the equilibrium mentioned above.     

东方田鼠警觉对其功能反应的作用格局

植食性哺乳动物功能反应描述了摄入率与植物可利用性变量的动态关系。动物警觉因占用了处理食物时间即觅食中断时间而延迟与下一口食物相遇,引致摄入率降低,进而对功能反应构型产生影响。在新鲜白三叶叶片构成的各类食物密集斑块上,测定东方田鼠觅食行为,建立功能反应函数模型,检验觅食中断对功能反应的作用格局及机制。结果发现,除小叶片斑块觅食中断时间在觅食活动中所占的比例较低外,在大、中型叶片斑块的觅食中断时间比例均达到15.42%-26.82%;尽管,觅食中断使摄入率降低了33%,但东方田鼠功能反应仍为Ⅱ型功能反应。除东方田鼠采食时间及觅食回合时间随叶片重增大保持相对稳定外,处理时间及觅食中断时间均随叶片重增大呈线性递增趋势;采食时间、处理时间及觅食中断时间随口量的增大呈线性递增趋势;采食率随叶片重和口量增大呈指数递减趋势。研究结果揭示,东方田鼠因警觉引起的觅食中断事件是导致采食率及摄入率降低的主要因子。摄入率测定值与模型预测值的线性回归极显著(P < 0.01),表明,新建立的功能反应模型具有良好的可预测性。推测,东方田鼠因警觉引起采食率及摄入率减小的代价,是以延长觅食时间来补偿的。研究结果充分验证了,在可利用性植物密集斑块,由植物大小调控的动物口量决定其摄入率,且受采食和处理食物竞争及觅食中断的制约,其功能反应为Ⅱ型功能反应的假说。     

Measuring the functional responses of farmland birds: an example for a declining seed-feeding bunting

1. Many farmland bird species have undergone significant declines. It is important to predict the effect of agricultural change on these birds and their response to conservation measures. This requirement could be met by mechanistic models that predict population size from the optimal foraging behaviour and fates of individuals within populations. A key component of these models is the functional response, the relationship between food and competitor density and feeding rate. 2. This paper describes a method for measuring functional responses of farmland birds, and applies this method to a declining farmland bird, the corn bunting Miliaria calandra L. We derive five alternative models to predict the functional responses of farmland birds and parameterize these for corn bunting. We also assess the minimum sample sizes required to predict accurately the functional response. 3. We show that the functional response of corn bunting can be predicted accurately from a few behavioural parameters (searching rate, handling time, vigilance time) that are straightforward to measure in the field. These parameters can be measured more quickly than the alternative of measuring the functional response directly. 4. While corn bunting violated some of the assumptions of Holling's disk equation (model 1 in our study), it still provided the most accurate fit to the observed feeding rates while remaining the most statistically simple model tested. Our other models may be more applicable to other species, or corn bunting feeding in other locations. 5. Although further tests are required, our study shows how functional responses can be predicted, simplifying the development of mechanistic models of farmland bird populations.     

Modeling nonbreeding distributions of shorebirds and waterfowl in response to climate change

To identify areas on the landscape that may contribute to a robust network of conservation areas, we modeled the probabilities of occurrence of several en route migratory shorebirds and wintering waterfowl in the southern Great Plains of North America, including responses to changing climate. We predominantly used data from the eB ird citizen‐science project to model probabilities of occurrence relative to land‐use patterns, spatial distribution of wetlands, and climate. We projected models to potential future climate conditions using five representative general circulation models of the Coupled Model Intercomparison Project 5 (CMIP 5). We used Random Forests to model probabilities of occurrence and compared the time periods 1981–2010 (hindcast) and 2041–2070 (forecast) in “model space.” Projected changes in shorebird probabilities of occurrence varied with species‐specific general distribution pattern, migration distance, and spatial extent. Species using the western and northern portion of the study area exhibited the greatest likelihoods of decline, whereas species with more easterly occurrences, mostly long‐distance migrants, had the greatest projected increases in probability of occurrence. At an ecoregional extent, differences in probabilities of shorebird occurrence ranged from ?0.015 to 0.045 when averaged across climate models, with the largest increases occurring early in migration. Spatial shifts are predicted for several shorebird species. Probabilities of occurrence of wintering Mallards and Northern Pintail are predicted to increase by 0.046 and 0.061, respectively, with northward shifts projected for both species. When incorporated into partner land management decision tools, results at ecoregional extents can be used to identify wetland complexes with the greatest potential to support birds in the nonbreeding season under a wide range of future climate scenarios.     

Breeding biology of the Kentish Plover Charadrius alexandrinus in the Sabkhat Al-Fasl Lagoons,Saudi Arabia (Aves: Charadriiformes)

The breeding biology of the Kentish Plover Charadrius alexandrinus was studied in the Sabkhat Al-Fasl Lagoons of Saudi Arabia, where ground temperatures may ex- ceed 55°C in summer. Although halophytic bushes are abundant, this species seems to prefer nesting at exposed sites. Biparental brood care was common: the females were absent in only three out of 24 families. Kentish Plovers attended their nests more than 80% over the full day and more than 90% of the time during day-time, and the number of change-overs increased during the hottest parts of the day which could be due to the possibility that a single parent cannot protect the eggs and itself from overheating.     

Harvest rates and foraging strategies in Negev Desert gerbils

We examined the foraging strategy and quantified the foragingtraits of two nocturnal rodent species, Allenby's gerbil (Gerbillusallenbyi) and the greater Egyptian sand gerbil (Gerbillus pyramidum).In the laboratory, both species used two distinct foragingstrategies: either they immediately consumed seeds found ina patch (seed tray); or they collected and delivered the seedsto their nest box for later consumption. Moreover, we founda transition in foraging strategy among individual G. allenbyi under laboratory conditions; they all began by consuming theseeds on the tray and, after 7 days on average, switched tothe collecting strategy. By contrast, in the field both speciesused only one foraging strategy; they collected and deliveredthe seeds to their burrow or to surface caches for later consumption.Furthermore, G. allenbyi and G. pyramidum collected seeds atsignificantly higher rates in the field than in the laboratorybecause the seed encounter rates for both species were higherin the field. This suggests that in natural conditions, probablyinvolving predation risk and competitive pressure, gerbilsmust respond in two ways: (1) they must choose a foraging strategythat reduces predation risk by minimizing time spent feedingoutside their burrows; and (2) they must forage more efficiently.In the field, seed handling time of the larger species, G. pyramidum, was shorter than that of the smaller one, G. allenbyi.This difference may give G. pyramidum an advantage when resourcelevels are high and when most of a forager's time is spent handling seeds rather than searching for more seeds. Additionally,our field study showed that the seed encounter rate of G. allenbyiwas higher than that of G. pyramidum. This difference may giveG. allenbyi an advantage when resource levels are low and whensearching occupies most of the forager's time. The differentadvantages that each species has over the other, under differentconditions, may well be factors promoting their coexistenceover a wide range of resource densities.     

根田鼠几种觅食功能反应模型的比较

植物组织空间排列对植食性哺乳动物功能反应的作用,是觅食生态学的热点问题之一。以新鲜紫花苜蓿叶片为食物,改变苜蓿叶片大小调控根田鼠口量,改变叶片间距调控叶片密度,设置叶片空间异质性斑块。在空间异质性斑块上测定根田鼠的觅食行为及其参数,检验植食性哺乳动物4种功能反应模型的预测性。除复合模型参数Rmax、h和Vmax及口量模型参数Rmax的最大似然估计值与测定值近似外,其它模型参数的估计值与测定值均存在较大差异。根田鼠摄入率测定值与4种模型预测值的线性回归均显著(P0.01),但与复合模型预测值的线性拟合效果最佳。表明,复合模型能很好地解释根田鼠觅食与行走的竞争对摄入率调节的动态。根田鼠复合模型存在调节其功能反应机制转变的距离临界值(d)。叶片间距大于该值时,叶片密度调节摄入率;叶片间距小于该值时,口量调节摄入率。结果充分地验证了提出的特定假设:在植物密集条件下,植物大小能调节植食性小型哺乳动物根田鼠的摄入率;在植物稀疏条件下,植物密度调节其摄入率。     

Characterization of 36 polymorphic microsatellite loci in the Kentish plover (Charadrius alexandrinus) including two sex‐linked loci and their amplification in four other Charadrius species

We isolated 45 new Kentish plover (Charadrius alexandrinus) microsatellite loci. These were tested for polymorphism in 42 Kentish plovers breeding in the Çukurova Delta, Turkey. Thirty‐six of the 45 loci were polymorphic with observed heterozygosity varying between 0.22 and 0.93. Genotypes of individuals of known sex indicated that two loci were sex‐linked (Calex‐26 is located on the Z chromosome and Calex‐31 on the W chromosome). Additionally, we tested all loci for amplification in four other species of Charadridae (Kittlitz's plover, Madagascar plover, three‐banded plover and white‐fronted plover). On average 34 loci amplified per species (range 29–36).     

植食性哺乳动物觅食功能反应模型机制的检验

植食性哺乳动物在食物密集斑块的觅食为 型功能反应。在新鲜苜蓿叶片构成的食物密集斑块上 ,以高原鼠兔作为实验动物 ,检验植食性哺乳动物的觅食功能反应及其模型机制。食物大小可调节高原鼠兔的口量 ,尽而控制其瞬时摄入率 ;高原鼠兔觅食叶片的口量 S与瞬时摄入率 I存在渐近的函数关系 ,为 型功能反应 ;高原鼠兔的食物收获率 B随口量 S的增加呈非线性递减 ;最大处理速率 Rmax的测定值与模型的预测值极为近似 ;瞬时摄入率 I的测定值与模型的预测值线性回归显著 ( P<0 .0 1 )。研究结果充分验证了提出的假设 :植食性哺乳动物 型功能反应模型能有效地预测其摄入率的动态 ;植食性哺乳动物收获与咀嚼间的竞争能调其收获率和摄入率     

Evolution of prey behavior in response to changes in predation regime: damselflies in fish and dragonfly lakes

In a large behavioral experiment we reconstructed the evolution of behavioral responses to predators to explore how interactions with predators have shaped the evolution of their prey's behavior. All Enallagma damselfly species reduced both movement and feeding in the presence of coexisting predators. Some Enallagma species inhabit water bodies with both fish and dragonflies, and these species responded to the presence of both predators, whereas other Enallagma species inhabit water bodies that have only large dragonflies as predators, and these species only responded to the presence of dragonflies. Lineages that shifted to live with large dragonflies showed no evolution in behaviors expressed in the presence of dragonflies, but they evolved greater movement in the absence of predators and greater movement and feeding in the presence of fish. These results suggest that Enallagma species have evolutionarily lost the ability to recognize fish as a predator. Because species coexisting with only dragonfly predators have also evolved the ability to escape attacking dragonfly predators by swimming, the decreased predation risk associated with foraging appears to have shifted the balance of the foraging/predation risk trade-off to allow increased activity in the absence of mortality threats to evolve in these lineages. Our results suggest that evolution in response to changes in predation regime may have greater consequences for characters expressed in the absence of mortality threats because of how the balance between the conflicting demands of growth and predation risk are altered.     

Spatial and temporal fluctuations in presence and use of chironomid prey by shorebirds in the Odiel saltpans, south-west Spain

We studied the seasonal variation in abundance and distribution of shorebirds and chironomid Chironomus salinarius larvae in both traditional and industrial salines in the Odiel marshes, south-west Spain, in 2001. We selected 12 ponds that were representative of the different phases of the salt production process. The benthic chironomids were sampled in each pond every 2 months, and the birds were counted weekly. Chironomid larvae were most abundant in spring and autumn, and in the ponds of lower salinity. The density of larvae averaged 7023 ± 392 m⁻² (±SE) over the six sampling events. Shorebirds were always more abundant at high tide than at low tide, and were especially abundant during the spring and autumn migration periods when up to 20,775 birds were counted. A total of 24 species were recorded, six of which were present in internationally important numbers. The salines were especially important as foraging and roosting habitat during migration. The percentage of birds that were feeding in the ponds was positively correlated with the abundance of chironomid larvae at accessible depths. The number of feeding birds was also higher in ponds with more chironomid larvae available. Despite more intensive management, industrial salines held higher densities of birds and a similar abundance of chironomids when compared with traditional salines.     

Preference in patchy landscapes: the influence of scale-specific intake rates and variance in reward

Understanding the responses of foragers to patchy distributionsof resources has formed a fundamental challenge in behavioralecology. Two currencies have been used to assess the patch preferencesof herbivores—intake rate maximization and risk sensitivity.We wished to understand if small mammalian foragers, collaredlemmings (Dichrostonyx groenlandicus), choose patches to maximizefood intake rate or to reduce risk of starvation in "variable"environments. Moreover, we examined the possibility that maximizingintake rate depends on the spatial scale of patchiness. We designedan experiment offering two alternative patches of food, varyingthe predictability of food rewards and the "potential intakerate" at different spatial scales. Collared lemmings did notconsistently select patches that maximized their intake rateat either scale studied. Instead, they chose patches offeringthe least variation in food reward over the course of the experiment.Collared lemmings used prior knowledge gained from previousforaging bouts to assess food variability. We interpret theseresults as evidence for risk-averse foraging strategies, whichare predicted for continuous foragers aiming to minimize riskof starvation.     

Noise-induced reduction in the attack rate of a planktivorous freshwater fish revealed by functional response analysis

1. Anthropogenic noise can affect animals physically, physiologically, and behaviourally. Although individual responses to noise are well documented, the consequences in terms of community structure, species coexistence, and ecosystem functioning remain fairly unknown.
2. The impact of noise on predation has received a growing interest and alterations in trophic links are observed when animals shift from foraging to stress-related behaviours, are distracted by noise, or because of acoustic masking. However, the experimental procedures classically used to quantify predation do not inform on the potential demographic impact on prey.
3. We derived the relationship between resource use and availability (the functional response) for European minnows (Phoxinus phoxinus) feeding on dipteran larvae (Chaoborus sp.) under two noise conditions: ambient noise and ambient noise supplemented with motorboat noise. The shape and magnitude of the functional response are powerful indicators of population outcomes and predator–prey dynamics. We also recorded fish behaviour to explore some proximate determinants of altered predation.
4. For both noise conditions, fish displayed a saturating (type II) functional response whose shape depends on two parameters: attack rate and handling time. Boat noise did not affect handling time but significantly reduced attack rate, resulting in a functional response curve of the same height but with a less steep initial slope. Fish exhibited a stress-related response to noise including increased swimming distance, more social interactions, and altered spatial distribution.
5. Our study shows the usefulness of the functional response approach to study the ecological impacts of noise and illustrates how the behavioural responses of predators to noise can modify the demographic pressure on prey. It also suggests that prey availability might mediate the negative effect of noise on predation. Community outcomes are expected if the reduced consumption of the main food sources goes with the overconsumption of alternative food sources, changing the distribution pattern of interaction strengths. Predation release could also trigger a trophic cascade, propagating the effect of noise to lower trophic levels.

     

Temperature dependence of the functional response

The Arrhenius equation has emerged as the favoured model for describing the temperature dependence of consumption in predator-prey models. To examine the relevance of this equation, we undertook a meta-analysis of published relationships between functional response parameters and temperature. We show that, when plotted in lin-log space, temperature dependence of both attack rate and maximal ingestion rate exhibits a hump-shaped relationship and not a linear one as predicted by the Arrhenius equation. The relationship remains significantly downward concave even when data from temperatures above the peak of the hump are discarded. Temperature dependence is stronger for attack rate than for maximal ingestion rate, but the thermal optima are not different. We conclude that the use of the Arrhenius equation to describe consumption in predator-prey models requires the assumption that temperatures above thermal optima are unimportant for population and community dynamics, an assumption that is untenable given the available data.     

Satiation and the functional response: a test of a new model

Abstract. 1. A model of the functional response to prey density is derived to include the reduction in time available for search, T_s , resulting from predator satiation.
2. For larger prey items predator satiation occurs at each prey capture and T_s is reduced by the attack time and digestive pause of a series of attack cycles. For small prey items predator foraging is continuous at low densities with T_s reduced solely by attack time. At higher densities predator satiation occurs after the capture of several small prey items and T_s is reduced by the attack time and digestive pause of a series of foraging cycles.
3. A comparison of the predicted asymptotic level of prey capture using experimentally estimated parameter values, with the maximum consumption of aphids by larval and adult coccinellids provides a test of the satiation model.
4. The limitation of prey capture by predator satiation is discussed with reference to handling time and the success of coccinellids in biological control.