The evolution of müllerian mimicry in multispecies communities

Prey species that are unprofitable to attack often share conspicuous colours and patterns with other coexisting defended species. This phenomenon, termed müllerian mimicry, has long been explained as a consequence of selection on defended prey to adopt a common way of advertising their unprofitability. However, studies using two unpalatable prey types have not always supported this theory. Here we show, using a system of humans hunting for computer-generated prey, that predators do not always generate strong selection for mimicry when there are two unprofitable prey types. By contrast, we demonstrate that when predators are faced with a range of different prey species, selection on unprofitable prey to resemble one another can be intense. Here the primary selective force is not one in which predators evaluate the profitabilities of distinct prey types independently, but one in which predators learn better to avoid unprofitable phenotypes that share traits distinguishing them from profitable prey. This need to simplify decision making readily facilitates the spread of imperfect mimetic forms from rarity, and suggests that müllerian mimicry is more likely to arise in multispecies communities.     

The evolution of inaccurate mimics

Instances of strikingly accurate Batesian mimicry (in which a palatable prey organism closely resembles an aversive model) are often cited to illustrate the power of natural selection. Less attention has been paid to those mimics, such as many hoverfly (Syrphidae) mimics of wasps or bees, that resemble their models only poorly. Attempts to provide an adaptive explanation for imperfect mimicry have suggested that what seems a crude resemblance to human observers may appear a close match to predators, or that inaccurate mimics may bear a general resemblance to several different model species. I show here, however, that truly inaccurate mimicry of a single model organism may be favoured over perfect resemblance, by kin selection. Signal detection theory predicts that predators will modify their level of discrimination adaptively in response to the relative frequencies and similarity of models and mimics. If models are rare and/or weakly aversive, greater local similarity of mimics can thus lead to greater attack rates. Where individual mimics are related to others in their vicinity, kin selection will then oppose the evolution of accurate mimicry.     

Selection overrides gene flow to break down maladaptive mimicry

Predators typically avoid dangerous species, and batesian mimicry evolves when a palatable species (the 'mimic') co-opts a warning signal from a dangerous species (the 'model') and thereby deceives its potential predators. Because predators would not be under selection to avoid the model and any of its look-alikes in areas where the model is absent (that is, allopatry), batesian mimics should occur only in sympatry with their model. However, contrary to this expectation, batesian mimics often occur in allopatry. Here we focus on one such example--a coral snake mimic. Using indirect DNA-based methods, we provide evidence suggesting that mimics migrate from sympatry, where mimicry is favoured, to allopatry, where it is disfavoured. Such gene flow is much stronger in nuclear genes than in maternally inherited mitochondrial genes, indicating that dispersal by males may explain the presence of mimetic phenotypes in allopatry. Despite this gene flow, however, individuals from allopatry resemble the model less than do individuals from sympatry. We show that this breakdown of mimicry probably reflects predator-mediated selection acting against individuals expressing the more conspicuous mimetic phenotype in allopatry. Thus, although gene flow may explain why batesian mimics occur in allopatry, natural selection may often override such gene flow and promote the evolution of non-mimetic phenotypes in such areas.     

Predator learning favours mimicry of a less-toxic model in poison frogs

Batesian mimicry--resemblance of a toxic model by an edible mimic--depends on deceiving predators. Mimetic advantage is considered to be dependent on frequency because an increase in mimic abundance leads to breakdown of the warning signal. Where multiple toxic species are available, batesian polymorphism is predicted--that is, mimics diversify to match sympatric models. Despite the prevalence of batesian mimicry in nature, batesian polymorphism is relatively rare. Here we explore a poison-frog mimicry complex comprising two parapatric models and a geographically dimorphic mimic that shows monomorphism where models co-occur. Contrary to classical predictions, our toxicity assays, field observations and spectral reflectances show that mimics resemble the less-toxic and less-abundant model. We examine "stimulus generalization" as a mechanism for this non-intuitive result with learning experiments using naive avian predators and live poison frogs. We find that predators differ in avoidance generalization depending on toxicity of the model, conferring greater protection to mimics resembling the less-toxic model owing to overlap of generalized avoidance curves. Our work supports a mechanism of toxicity-dependent stimulus generalization, revealing an additional solution for batesian mimicry where multiple models coexist.     

A comparative analysis of the evolution of imperfect mimicry

Although exceptional examples of adaptation are frequently celebrated, some outcomes of natural selection seem far from perfect. For example, many hoverflies (Diptera: Syrphidae) are harmless (Batesian) mimics of stinging Hymenoptera. However, although some hoverfly species are considered excellent mimics, other species bear only a superficial resemblance to their models and it is unclear why this is so. To evaluate hypotheses that have been put forward to explain interspecific variation in the mimetic fidelity of Palearctic Syrphidae we use a comparative approach. We show that the most plausible explanation is that predators impose less selection for mimetic fidelity on smaller hoverfly species because they are less profitable prey items. In particular, our findings, in combination with previous results, allow us to reject several key hypotheses for imperfect mimicry: first, human ratings of mimetic fidelity are positively correlated with both morphometric measures and avian rankings, indicating that variation in mimetic fidelity is not simply an illusion based on human perception; second, no species of syrphid maps out in multidimensional space as being intermediate in appearance between several different hymenopteran model species, as the multimodel hypothesis requires; and third, we find no evidence for a negative relationship between mimetic fidelity and abundance, which calls into question the kin-selection hypothesis. By contrast, a strong positive relationship between mimetic fidelity and body size supports the relaxed-selection hypothesis, suggesting that reduced predation pressure on less profitable prey species limits the selection for mimetic perfection.     

Multiple benefits of gregariousness cover detectability costs in aposematic aggregations

Understanding the early evolution of aposematic (warning) coloration has been a challenge for scientists, as a new conspicuous morph in a population of cryptic insects would have a high predation risk and would probably die out before local predators learnt to avoid it. Fisher presented the idea of aggregation benefit through the survival of related individuals; however, his theory has been strongly debated as the mechanisms that favour grouping have never been explored experimentally with the incorporation of detectability costs. Here we create a comprehensive 'novel world' experiment with the great tit (Parus major) as a predator to explore simultaneously the predation-related benefits and costs for aposematic aggregated prey, manipulating both group size and signal strength. Our results show that grouping would have been highly beneficial for the first aposematic prey individuals surrounded by naive predators, because (1) detectability risk increased only asymptotically with group size; (2) additional detectability costs due to conspicuous signals were marginal in groups; (3) even naive predators deserted the group after detecting unpalatability (dilution effect); and (4) avoidance learning of signal was faster in groups. None of these mechanisms require kin selection.     

Three-butterfly system provides a field test of müllerian mimicry

In 1879, Müller proposed that two brightly coloured distasteful butterfly species (co-models) that share a single warning-colour pattern would benefit by spreading the selective burden of educating predators. The mutual benefit of sharing warning signals among distasteful species, so-called müllerian mimicry, is supported by comparative evidence, theoretical studies and laboratory simulations; however, to date, this key exemplar of adaptive evolution has not been experimentally tested in the field. To measure natural selection generated by müllerian mimicry, I exploited the unusual polymorphism of Heliconius cydno (Lepidoptera: Nymphalidae). Here I show increased survival of H. cydno morphs that match locally abundant monomorphic co-model species. This study demonstrates müllerian mimicry in the field. It also shows that müllerian mimicry with several co-models generates geographically divergent selection, which explains the existence of polymorphism in distasteful species with warning coloration.     

Animal mimicry: choosing when to be a cleaner-fish mimic

Mimicry in vertebrates is usually a permanent state--mimics resemble and normally accompany their model throughout the life stages during which they act as mimics. Here we show that the bluestriped fangblenny fish (Plagiotremus rhinorhynchos), which aggressively attacks other coral-reef fish, can turn off the mimetic colours that disguise it as the benign bluestreak cleaner wrasse, Labroides dimidiatus, and assume a radically different appearance. This opportunistic facultative mimicry extends the fangblenny's scope by allowing it to blend into shoals of small reef fish as well as to remain inconspicuous at cleaning stations.     

An obligate brood parasite trapped in the intraspecific arms race of its hosts

Reciprocal selection pressures often lead to close and adaptive matching of traits in coevolved species. A failure of one species to match the evolutionary trajectories of another is often attributed to evolutionary lags or to differing selection pressures across a geographic mosaic. Here we show that mismatches in adaptation of interacting species--an obligate brood parasitic duck and each of its two main hosts--are best explained by the evolutionary dynamics within the host species. Rejection of the brood parasite's eggs was common by both hosts, despite a lack of detectable cost of parasitism to the hosts. Egg rejection markedly reduced parasite fitness, but egg mimicry experiments revealed no phenotypic natural selection for more mimetic parasitic eggs. These paradoxical results were resolved by the discovery of intraspecific brood parasitism and conspecific egg rejection within the hosts themselves. The apparent arms race between species seems instead to be an incidental by-product of within-species conflict, with little recourse for evolutionary response by the parasite.     

The effects of competition and predation on diversification in a model adaptive radiation

Much of life's diversity is thought to have arisen through successive rounds of adaptive radiation-the rapid diversification of a lineage into a range of ecologically and phenotypically distinct species. Both resource competition and predation have been suggested as mechanisms driving this process, although the former is better studied than the latter. Here we show experimentally how predation by a protist, Tetrahymena thermophila, affects diversification in a model adaptive radiation of the bacterial prey, Pseudomonas fluorescens. We estimate the frequency-dependent fitness functions of competing niche-specialist prey in the presence and absence of predation, and use these to test hypotheses about the extent (measured as the number of new genotypes) and rate of diversification. Competition and predation independently generated diversifying selection that we show is capable of driving prey diversification to similar extents but at different rates, diversification being markedly delayed in the presence of predators. The cause of this delay stems from weaker diversifying selection due to the reduction in prey density caused by predation. Our results suggest that predation may play an under-appreciated role in driving adaptive radiations.     

Coexistence of two sympatric and mOrpho1ogically similar bat species Rhinolophus affnis and Rhinolophus pearsoni

Ecologists have long focused on the coexistence of sympatric species.Here,we investigated two horseshoe bat species,Rhinolophus affinis and Rhinolophus pearsoni inhabited in the same cave,for their foraging strategies,niche differentiation,prey selection,and their coexistence status.These two species of horseshoe bats were ditierent in the dominant frequency of their echolocation calls.but similar in their morphology.We found evidence for prey selectivity although there was a high degree of overlap in prey categories and sizes.R affinis and R pearsoni foraged on 16 and 7 categories insects,respectively,with Pyralidae,Geometridae,Melolonthidae dominating their diets.The degree of trophic niche overlap was 0.69.Pairwise comparisons suggested that there was no obvious differentiation in prey categories and size.However,high prey availability in the environment(Simpson diversity index=0.79 and Margalef richness index=4.121 contributed to their coexistence by dampening the interspecific competition.Since there are one or more mechanisms facilitating species coexistence in a community,our results suggest that the spatial niche differentiation in foraging microhabitats and in foraging habitats at landscape scale may promote the coexistence of the two bat species.However,additional field data are needed to confirm this speculation.     

Coexistence of two sympatric and morphologically similar bat species Rhinolophus affinis and Rhinolophus pearsoni

Ecologists have long focused on the coexistence of sympatric species. Here, we investigated two horseshoe bat species, Rhinolophus affinis and Rhinolophus pearsoni inhabited in the same cave, for their foraging strategies, niche differentiation, prey selection, and their coexistence status. These two species of horseshoe bats were different in the dominant frequency of their echolocation calls, but similar in their morphology. We found evidence for prey selectivity although there was a high degree of overlap in prey categories and sizes. R. affinis and R. pearsoni foraged on 16 and 7 categories insects, respectively, with Pyralidae, Geometridae, Melolonthidae dominating their diets. The degree of trophic niche overlap was 0.69. Pairwise comparisons suggested that there was no obvious differentiation in prey categories and size. However, high prey availability in the environment （Simpson diversity index = 0.79 and Margalef richness index = 4.12） contributed to their coexistence by dampening the interspecific competition. Since there are one or more mechanisms facil- itating species coexistence in a community, our results suggest that the spatial niche differentiation in foraging rnicrohabitats and in foraging habitats at landscape scale may promote the coexistence of the two bat species. However, additional field data are needed to confirm this speculation.     

福建沿海地区农业生态环境脆弱度模糊综合评判

根据福建沿海地区农业脆弱生态环境的特点,以人均水资源、森林覆盖率、人均耕地面积、水土流失、干燥指数、农药使用量、化肥使用量、人均GDP、农民人均收入、恩格尔系数、人口密度、文盲率、高等教育率等作为评价指标,利用层次分析法分配各指标权重,通过模糊综合评判模型运算和隶属度分析,将福建沿海地区农业脆弱生态环境分为较严重脆弱、中等脆弱、一般脆弱、无脆弱4个等级.研究结果为该地区农业环境可持续发展提供科学依据.     

Coexistence of two sympatric and morphologically similar bat species Rhinolophus affinis and Rhinolophus pearsoni

Ecologists have long focused on the coexistence of sympatric species. Here, we investigated two horseshoe bat species, Rhinolophus affinis and R. pearsoni inhabited in the same cave, for their foraging strategies, niche differentiation, prey selection, and their coexistence status. These two species of horseshoe bats were different in the dominant frequency of their echolocation calls but similar in their morphology. We found evidence for prey selectivity although there was a high degree of overlap in prey categories and sizes. R. affinis and R. pearsoni foraged on 16 and 7 categories insects, respectively, with Pyralidae, Geometridae, Melolonthidae dominating their diets. The degree of trophic niche overlap was 0.69. Pairwise comparisons suggested that there was no obvious differentiation in prey categories and size. However, high prey availability in the environment (Simpson diversity index = 0.79 and Margalef richness index ＝ 4.12) contributed to their coexistence by dampening the interspecific competition. Since there are one or more mechanisms facilitating species coexistence in a community, our results suggest that the spatial niche differentiation in foraging microhabitats and in foraging habitats at landscape scale may promote the coexistence of the two bat species. However, additional field data are needed to confirm this speculation.     

Parasitoid secretions provoke ant warfare

Insect social parasites are extreme specialists that typically use mimicry or stealth to enter ant colonies to exploit the rich, but fiercely protected, resources within their nests. Here we show how a parasitic wasp (parasitoid) contrives to reach its host, itself an endangered species of social parasite that lives inside the brood chambers of ant nests, by releasing semiochemicals to induce in-fighting between worker ants, locking the colony in combat and leaving it underprotected. Four of these chemicals are new to biology and have the potential to control pest species by inducing different agonistic behaviours in ants.     

Competition and phylogeny determine community structure in Müllerian co-mimics

Until recently, the study of negative and antagonistic interactions (for example, competition and predation) has dominated our understanding of community structure, maintenance and assembly. Nevertheless, a recent theoretical model suggests that positive interactions (for example, mutualisms) may counterbalance competition, facilitating long-term coexistence even among ecologically undifferentiated species. Müllerian mimics are mutualists that share the costs of predator education and are therefore ideally suited for the investigation of positive and negative interactions in community dynamics. The sole empirical test of this model in a Müllerian mimetic community supports the prediction that positive interactions outweigh the negative effects of spatial overlap (without quantifying resource acquisition). Understanding the role of trophic niche partitioning in facilitating the evolution and stability of Müllerian mimetic communities is now of critical importance, but has yet to be formally investigated. Here we show that resource partitioning and phylogeny determine community structure and outweigh the positive effects of Müllerian mimicry in a species-rich group of neotropical catfishes. From multiple, independent reproductively isolated allopatric communities displaying convergently evolved colour patterns, 92% consist of species that do not compete for resources. Significant differences in phylogenetically conserved traits (snout morphology and body size) were consistently linked to trait-specific resource acquisition. Thus, we report the first evidence, to our knowledge, that competition for trophic resources and phylogeny are pivotal factors in the stable evolution of Müllerian mimicry rings. More generally, our work demonstrates that competition for resources is likely to have a dominant role in the structuring of communities that are simultaneously subject to the effects of both positive and negative interactions.     

山东省经济发展与环境污染关系实证研究

环境库兹涅兹曲线(EKC)是描述污染问题与经济发展之间关系的常用模型,一般情况下呈倒U形.通过对山东省1990～2005年的人均GDP与工业三废排放量之间的关系分析,分别建立二次和三次环境库兹涅兹曲线模型.研究发现,山东省人均废气排放量与人均GDP符合倒U型模型,而人均二氧化硫排放量、人均烟尘排放量、人均固体废弃物排放量与人均GDP之间不符合倒U型模型.文章从环境保护成效、环境政策成本和经济增长模式三个方面评述了山东省环境经济政策,并提出环境保护与经济发展综合决策机制和政策框架.     

人口红利、财政支出与经济增长

运用描述统计、稳定性检验、三段线性回归模型和VAR模型分析财政支出占GDP比重随时间的变动趋势、人均财政支出和人均GDP的相互关系和影响人均财政支出因素的动态变化。结果表明,20世纪70年代以来,我国实行的计划生育政策促进了人均财政支出与人均GDP的长期稳定增长;1981—2011年,人均财政支出、人均GDP、人均固定资产投资和人均工业总产值之间存在长期稳定的均衡关系。     

济南市工业"三废"排放的环境库兹涅茨曲线特征分析

选取济南市1994-2003年经济与环境数据,建立单位GDP污染排放量模型和人均GDP污染排放量模型,分析工业“三废”与人均GDP增长的环境库兹涅茨曲线(EKC)。从模型模拟结果看,单位GDP污染排放量总体呈现下降趋势,表明济南市产业结构在十几年的演变过程中发生了明显的变化。发现济南市经济发展与环境污染水平的关系总体上符合EKC特征;济南市经济目前仍处于工业化进程的中期阶段,环境污染物排放量EKC的转折点尚未达到。     

基于灰色马氏链模型的农村居民人均收入预测

对农村居民人均收入变化趋势进行研究,将灰色模型与马氏链模型相结合,建立灰色马氏链预测模型,对河南省新乡市农村居民人均收入进行预测.与灰色预测模型相比,灰色马氏链模型能够提高预测精度.利用灰色马氏链模型对2017—2025年新乡市农村居民人均收入进行了预测,结果表明,该市农村居民人均收入有明显的上升趋势.