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2018年5月26日托福阅读真题解析2018年5月26日的托福阅读考试终于结束了，同学们是不是非常的兴奋，是不是想了解自己考得怎么样呢？下面就和店铺一起来看看2018年5月26日托福阅读真题解析。
Passage One学科分类：社会题目：Two forces of urban growth内容回忆：首段：首先说城市增长有很多原因，比如经济增长、交通发展、科技进步。
然后除了这些之外引入了两个概念：centrifugal（分散化）和centripetal（中心化）。
（分类讨论）然后介绍manufact uring的发展对于城镇发展的促进作用：蒸汽机解放了产业对于流水的依赖性，早期的制造商人把工厂建造在城镇中心，主要有两个原因：1、城镇有丰富的劳动力，可以解决工厂劳动力来源；2、当时基本交通方式是以步行为主，所以要把工厂建立在人们步行可以达到的地方，这种布置促进了城镇中心化发展。
（第一题是个时间取反的推理题，问蒸汽机发明前制造业怎么样。
）第二段：虽然制造业对于商业发展有促进作用，但是随着社会的进步，经济的发展，零售业和其他行业的发展也集聚在城市中心，这使得城市的地价进一步提升，对于制造业来说成本就越来越高，加上铁路等运输方式的发展，制造业可以分布在城市郊区，这样降低成本，同时只需要修建较短的运输方式就可以和城市中心相连，所以城市开始扩张，制造业分布在市郊。
第三段：对于零售业的发展，零售业必须建在有大量居民的地区，同时因为可以在商店内陈列大量商品，所以零售业的利润一直都很高。
同时混凝建筑的发展、电梯的使用，促进了零售业的垂直分布，可以容纳不断扩张的产品和工作人员。
（出了一个句子简化题）第四段：交通运输业的发展促进了销售和产品生产的分离，人们可以无需面对面沟通，通过电话、电报沟通即可，进一步促进了城市的扩张。
Commercial office可以在市中心，接近人群和各种服务，同时也可以控制生产中心。
词汇题：Respectively=particularlySufficient=enoughProximity=in short distancesThus=in this wayPassage Two学科分类：动物学题目：Storage strategies of squirrels内容回忆：首段：有很多动物有储藏策略，但是松鼠的储藏手段在所有动物中尤其突出。
大学英语六级模拟试卷376(题后含答案及解析)题型有：1. Writing 2. Reading Comprehension (Skimming and Scanning) 3. Listening Comprehension 4. Reading Comprehension (Reading in Depth) 6. Error Correction 8. TranslationPart I Writing (30 minutes)1．Directions: For this part, you are allowed 30 minutes to write a composition on the topic: Precious Water. You should write at least 150 words following the outline given below:1.举例说明水对人类的重要性2. 举例说明我国所面临的水资源问题3. 为了生存和发展人们要……Precious Water正确答案：Precious Water No one doubts the importance of water. Human beings live on water, animals live on water, plants live on water and all living things live on water. You can’t imagine what would happen if there were no water in the world! China is one of the countries in the world that are badly short of water. Water problem has become more acute in recent years with the increase of water consumption and pollution. In big cities, the water problem has become even worse. With the growth of population, more and more water is needed. With the development of the industry, a large quantity of water has been and is being polluted. Water pollution is the most serious problem that China is facing. It has spread to rivers, lakes, and even the oceans. In order to survive, man has to be wise enough to treasure water and try m prevent it from being polluted. Man should also know that all resources in the world are limited, including water. Make good use of them, otherwise, man will be punished by nature. To treasure water is to treasure life: to protect our environment is to protect ourselves.Part II Reading Comprehension (Skimming and Scanning) (15 minutes)Directions: In this part, you will have 15 minutes to go over the passage quickly and answer the questions attached to the passage. For questions 1-4, mark：Y (for YES) if the statement agrees with the information given in the passage;N (for NO) if the statement contradicts the information given in the passage;NG (for NOT GIVEN)if the information is not given in the passage.Where Have All the People Gone? Germans are getting used to a new kind of immigrant. In 1998, a pack of wolves crossed the Neisse River on the Polish-German border. In the empty landscape of eastern Saxony, dotted with abandoned mines and declining villages, the wolves found plenty of deer and few humans. Five years later, a second pack split from the original, so there’re now two families of wolves in the region. A hundred years ago, a growing land-hungry population killed off the last of Germany’s wolves. Today, it’s the local humans whose numbers are under threat. Villages are empty, thanks to the region’s low birth rateand rural flight. Home to 22 of the world’s 25 lowest fertility rate countries, Europe will lose 30 million people by 2030, even with continued immigration. The biggest population decline will hit rural Europe. As Italians, Spaniards, Germans and others produce barely three-fifths of children needed to maintain status quo, and as rural flight sucks people into Europe’s suburbs and cities, the countryside will lose a quarter of its population. The implications of this demographic (人口的) change will be far-reaching.Environmental Changes The postcard view of Europe is of a continent where every scrap of land has long been farmed, fenced off and settled. But the continent of the future may look rather different. Big parts of Europe will renaturalize. Bears are back in Austria. In Swiss Alpine valleys, farms have been receding and forests are growing back. In parts of France and Germany, wildcats and wolves have re-established their ranges. The shrub and forest that grows on abandoned land might be good for deer and wolves, but is vastly less species-rich than traditional farming, with its pastures, ponds and hedges. Once shrub covers everything, you lose the meadow habitat. All the flowers, herbs, birds, and butterflies disappear. A new forest doesn’t get diverse until a couple of hundred years old. All this is not necessarily an environmentalist’s dream it might seem. Take the Greek village of Prastos. An ancient hill town, Prastos once had 1,000 residents, most of them working the land. Now only a dozen left, most in their 60s and 70s. The school has been closed since 1988. Sunday church bells no longer ting. Without farmers to tend the fields, rain has washed away the once fertile soil. As in much of Greece, land that has been orchards and pasture for some 2,000 years is now covered with dry shrub that, in summer, frequently catches fire.Varied Pictures of Rural Depopulation Rural depopulation is not new. Thousands of villages like Prastos dot Europe, the result of a century or more of emigration, industrialization, and agricultural mechanization. But this time it’s different because never has the rural birth rate so low. In the past, a farmer could usually find at least one of his offspring to take over the land. Today, the chances are that he has only a single son or daughter, usually working in the city and rarely willing to return. In Italy, more than 40% of the country’s 1.9 million farmers are at least 65 years old. Once they die out, many of their farms will join the 6 million hectares —one third of Italy’s farmland —that has already been abandoned. Rising economic pressures, especially from reduced government subsidies, will amplify the trend. One third of Europe’s farmland is marginal, from the cold northern plains to the dry Mediterranean (地中海) hills. Most of these farmers rely on EU subsides, since it’s cheaper to import food from abroad. Without subsidies, some of the most scenic European landscapes wouldn’t survive. In the Austrian or Swiss Alps, defined for centuries by orchards, cows, high mountain pastures, the steep valleys are labor-intensive to farm, with subsidies paying up to 90% of the cost. Across the border in France and Italy, subsidies have been reduced for mountain farming. Since then, across the southern Alps, villages have emptied and forests have grown back in. Outside the range of subsidies, in Bulgaria, Romania and Ukraine, big tracts of land are returning to wild.Big Challenges The truth is varied and interesting. While many rural regions of Europe are emptying out, others will experience something of a renaissance. Already, attractive areas within driving distance of prosperous cities areseeing robust revivals, driven by urban flight and an in-flooding of childless retirees. Contrast that with less-favored areas, from the Spanish interior to eastern Europe. These face dying villages, abandoned farms and changes in the land not seen for generations. Both types of regions will have to cope with steeply ageing population and its accompanying health and service needs. Rural Europe is the laboratory of demographic changes. For governments, the challenge has been to develop policies that slow the demographic decline or attract new residents. In some places such as Britain and France, large parts of countryside are reviving as increasingly wealthy urban middle class in search of second homes recolonises villages and farms. Villages in central Italy are counting on tourism to revive their town, turning farmhouses into hostels for tourists and hikers. But once baby boomers start dying out around 2020, populations will start to decline so sharply that there simply won’t be enough people to reinvent itself. It’s simply unclear how long current government policies can put off the inevitable. “We are now talking about civilized depopulation. We just have to make sure that old people we leave behind are taken care of.” Says Mats Johansson of Royal Institute of Technology in Stockholm. The biggest challenge is finding creative ways to keep up services for the rising proportion of seniors. When the Austrian village of Klans, thinly spread over the Alpine foothills, decided it could no longer afford a regular public bus service, the community set up a public taxi-on-demand service for the aged. In thinly populated Lapland where doctors are few and far between, tech-savvy Finns the rising demand for specialized health care with a service that uses videoconferencing and the Internet for remote medical examination. Another pioneer is the village of Aguaviva, one of rapidly depopulating areas in Spain. In 2000, Mayor Manznanares began offering free air-fares and housing for foreign families to settle in Aguaviva. Now the mud-brown town of about 600 has 130 Argentine and Romanian immigrants, and the town’s only school has 54 pupils. Immigration was one solution to the problem. But most foreign immigrants continue to prefer cities. And within Europe migration only exports the problem. Western European look towards eastern Europe as a source for migrants, yet those countries have ultra-low birth rates of their own. Now the increasingly worried European governments are developing policies to make people have more children, from better childcare to monthly stipends (津贴) linked to family size. But while these measures might raise the birth rate slightly, across the much of the ageing continent there are just too few potential parents around.2．The current rural depopulation in Europe is the result of long-term emigration and industrialization.A．YB．NC．NG正确答案：B解析：根据第二段可知，最近欧洲农村人口锐减主要是由于低出生率与农村人口迁往城市(the region’s low birth rate and rural flight)。
2020年大学英语六级考试强化：长难句翻译(1)1. The American economic system is organized around a basically private-enterprise, market-oriented economy in which consumers largely determine what shall be produced by spending their money in the marketplace for those goods and services that they want most.[参考译文]美国的经济是以基本的私有企业和市场导向经济为架构的，在这种经济中，消费者很大水准上通过在市场上为那些他们最想要的货品和服务付费来决定什么应该被制造出来。
2. Thus, in the American economic system it is the demand of individual consumers, coupled with the desire of businessmen to maximize profits and the desire of individuals to maximize their incomes that together determine what shall be produced and how resources are used to produce it.[参考译文]所以，在美国的经济体系中，个体消费者的需求与商人试图化其利润的欲望和个人想化其收入效用的欲望相结合，一起决定了什么应该被制造，以及资源如何被用来制造它们。
3. If, on the other hand, producing more of a commodity results in reducing its cost, this will tend to increase the supply offered by seller-producers, which in turn will lower the price and permit more consumers to buy the product.[参考译文]另一方面，如果大量制造某种商品导致其成本下降，那么这就有可能增加卖方和制造商能提供的供给，而这也就会反过来降低价格并允许更多的消费者购买产品。
2016年6月大学英语四级《仔细阅读》练习题(3) Agriculture is the number one industry in the United States and agricultural products are the country's leading export.American farmers manage to feed not only the total population of the United States,but also millions of other people throughout the rest of the world.Corn and soybean exports alone account for approximately75percent of the amount sold in world markets.This productivity,however,has its price.Intensive cultivation exposes the earth to the damaging forces of nature.Every year wind and water remove tons of rich soil from the nation's croplands.Each field is covered by a limited amount of topsoil,the upper layer of earth which is richest in the nutrients and minerals necessary for growing crops.Ever since the first farmers arrived in the Midwest almost200years ago，cultivation and,consequently, erosion have been decreasing the supply of topsoil.In the1830s,nearly two feet of rich,black top soil covered the Midwest.Today the average depth is only eight inches, and every decade another inch is blown or washed away.This erosion is steadily decreasing the productivity of valuable cropland.A United States Agricultural Department survey states that if erosion continues at its present rate,corn and soybean yields in the Midwest may drop as much as30percent over the next50years.So far,farmers have been able to compensate for the loss of fertile topsoil by applying more chemical fertilizers to their fields;however,while this practice hasincreased crop yields,it has been devastating for ecology.Agriculture has become one of the biggest polluters of the nation's precious water supply.Rivers,lakes,and underground reserves of water are being filled in and poisoned by soil and chemicals carried by drainage from eroding fields.Furthermore,fertilizers only replenish the soils they do not prevent its loss.26.The last sentence in the first paragraph gives an example to show_______.A.that American farmers manage to feed the total population of the U.S.B.the leading position of the U.S.farming in the worldC.how important American people consider their farmingD.that many people in the world rely on the export of the agricultural products of the U.S.A.27.In order to compensate for the loss of fertile topsoil,farmers have been_______.A.planting less corn and soybeanB.putting fertilizers on their fieldsC.preventing soil erosionD.decreasing the supply of top soil28.At the present rate,approximately how many years later the black top soil now covering the Midwest will completely be blown or washed away?A.120years later..B.80years later.C.50years later.D.100years later.29."This practice"in Paragraph4refers to_______.A.that farmers have lowered the yield of corn and soybeanB.that farmers have expanded croplandsC.that farmers have applied more chemical fertilizersD.that the top soil has been decreased greatly30.All of the following are statements about the disadvantages of fertilizers EXCEPT thatA.they replenish the soilB.they do not prevent the loss of soilC.they are destroying the ecologyD.they pollute the nation's water supply答案：26.D27.B28.B29.C30.A本文来源于中国大学网。
2021年6月英语四级阅读理解长难句精析【导语】梦想在前方，努力在路上。
对于考生来说，拿到证书就是我们向往的远方。
以下是“年月英语四级阅读理解长难句精析”，欢迎阅读参考！更多相关讯息请关注！【篇一】年月英语四级阅读理解长难句精析The reverse flow, from developed todeveloping countries, is on the rise,too.原文译文：从发达国家到发展中国家的逆向流动也有所增长。
四级词汇讲解：本句的主干是flow is on the rise。
from developedto developing countries为句子的插入语，起到对主语reverse flow的补充说明作用。
reverse flow意为“逆流”。
on the rise的意思是“在上升，在增长”：如：Dividends are on the rise, prices are on the rise and pressure will only continue to rise.分红在增加，价格在上升，而压力也只会继续增大。
英语四级考点归纳：插入语即句子中间插入的一部分，不充当任何句子成分，也不和句子成分发生结构关系，同时既不起连接作用，也不表示语气。
插入语大致分为以下几类:※用简短的句子结构作插入语，常见的有：I think, I hope,I guess,I believe,I suppose, I wonder, I tell you,I say, I'm afraid, I'sure,you see, you know, as youknow, that is, that is to say, what's more等，可以置于句中或句尾。
如：This diet,I think, will do good to your health.这类饮食，我感觉会有益于你的健康。
11月15日托福阅读真题（新东方版）第一篇考生回忆：本文属讲的是热带雨林树木的种类。
先讲了在温带地区树的种类非常好区分，但是在热带去不是这个样子。
接下来说开花的情况不一样。
在干湿分明的雨林中，花是随着雨季开放的。
在很多雨林一直都有雨，那么它们如何授粉?应该是一起同时开花。
接下来说叶子难以区分，大部分叶子要进化成很厚的样子，而且叶尖会滴水。
所以都长得很像。
另外一种辨别的方法是看树干。
虽然树干也很像。
接下来说在温带地区，树种没有那么难以辨别。
数量不会超过半打，但是热带的确是难以辨别。
第二篇考生回忆：主要讨论的是地球与宇宙年份的测定。
先说地球年份的测算。
一开始说可以通过陨石的年份来推测。
但是因为地球上面有水、火山和地壳运动，所以岩石的成分无法分析。
进一步的分析方法是岩石的辐射衰变。
得出来的结论是46亿年，这个测算的结果和月球上测算的结果是一样的。
因为月球上没有地球上的这些侵蚀，所以月球的测算数据相对准确。
接下来是宇宙年份的测算。
宇宙的年份重要是通过红移(red shift)来测量。
测算的结果发现宇宙一直在不断地膨胀。
第三篇考生回忆：讨论恐龙灭绝的原因。
首先说尝试解释恐龙灭绝的假说是在太多。
分析了6亿5千万年之前的底层，发现很好含钙的化石，其中就有恐龙的。
超过这个时间之后恐龙化石就再也找不到了，也就是认为恐龙灭绝了。
接下来提出一个假说，认为是辐射所导致恐龙基因变异导致恐龙灭绝。
恰好是每隔几百万年地球的南北磁极就会发生倒置，地磁场太弱，所以恐龙灭绝了。
后来又认为是陨石撞击地球，但是恐龙并没有立马就全部死绝，而是通过破坏全球的气候，通过影响底层生物链的生物最终使得恐龙饿死。
这个理论认为这些间接因素也很关键。
最后说火山运动也可能导致恐龙灭绝。
举了印度的例子，长时间的火山爆发导致了大片新火山岩的形成。
火山爆发喷出的灰烬遮盖了太阳，使得全球温带骤降。
但也有温室气体的排放会导致温度提高。
后来说这样的结果导致了新物种的大爆发。
雅思写作高分技巧：状语前置今天三立在线教育雅思网为大家带来的是雅思写作高分技巧：状语前置的相关资讯，备考的烤鸭们，赶紧来看看吧！状语前置就是把一个修饰动词的状语结构，如介词短语，分词形式或动词不定式引导的短语放到句首。
请看下面的句子：（1）Like self-awareness，this is also very difficult to achieve，but I think these are the two factors that may be the most important for achieving happiness.（2）Throughout the century，the largest quantity of water was used for agricultural purposes.（3）With a population of 176 million，the figures for Brazil indicate how high agricultural water consumption can be in some countries.使用状语前置的最大优点是让单调的句子有了跳跃的节奏感。
考官一天看上百张考卷，看到这样的句子也会心情愉悦。
注意：插入语此种语法结构是可以理解为是状语前置的另一种变体，它将状语结构提到了主句的主语和谓语之间。
插入语也是相对地道的英语表达方法。
请看以下几例：（1）Universities，when it is functioning well，should offer both theoretical knowledge as well as professional training.（2）So overall，I believe that，attending school from a young age is good for most children.插入语的功能和状语前置基本相似，都能使句子更有跳跃感和地道。
2008年12月一、Sustainable development is applied to just about everything from energy to clean water and economic growth, and as a result it has become difficult to question either the basic assumptions behind it or the way the concept is put to use.可持续发展适用于诸如能源，净水，经济增长等几乎所有的方面，因此，要对可持续发展的基本假设或其概念的实施方法提出质疑也变得日益困难。
This is especially true in agriculture, where sustainable development is often taken as the sole measure of progress without a proper appreciation of historical and cultural perspectives.这在农业方面尤其显著，可持续发展经常被认为是农业进步的唯一标准，而这并没有从历史和文化的角度进行适当的评估。
To start with, it is important to remember that the nature of agriculture has changed markedly throughout history, and will continue to do so .medieval agriculture in northern Europe fed, clothed and sheltered a predominantly rural society with a much lower population density than it is today.首先，重要的是认识到农业的本质随着历史的发展已经发生了显著的变化，并且这种变化仍将持续。
一、请将下面这段话翻译成英文：助人为乐，是中华民族优良传统之一。
通过“助人”，既向别人供给了帮助，又表达了一种自尊。
帮助他人要摈弃私心杂念，不能处处为个人利益着想。
遇事要多替别人考虑，主动伸手帮助那些需要帮助的人。
做到助人为乐，要偷快面对生活，不能自寻苦恼。
在帮助别人的同时，自己收获欢快，享受生活的乐趣。
做到助人为乐，要乐观行动起来，不能只说不做。
要脚踏实地(be down-to-earth), 热忱周到地为他人效劳，哪怕是简洁的小事，也要从一点一滴做起。
参考翻译：Being ready to help others is one of the fine traditions of Chinese nation.By helping others，one not only offered help to others,but also expressed one kind of self-respect.To help others,one should give up selfishness and shouldn”t consider his own interest all the time.Think more of others and initiatively give a hand to those that need help.To be ready to help others,one should live happily and avoid asking for trouble.When helping others, one can get happiness at the same time and enjoy the pleasure of life.To be ready to help others,one should take action actively instead of just saying it.Be down-to-earth, and offer service to others with passion.Even for the simple things,just start doing them bit by bit.翻译讲解：1.表达了一种自尊:可译为express one kind of self-respect。
INTRODUCTIONThroughout E urope, agricultural intensification, irriga-tion and yield improvement schemes, and abandonment of less-productive land, have caused serious declines of butterflies associated with traditionally managed humid meadows (Van Swaay & Warren, 1999; Maes & Van Dyck, 2001; Beneš et al., 2002). Such losses urge the col-lection of detailed ecological information on individual species that can be directly applied to practical conserva-tion measures (e.g., Neve et al., 1996; Fisher et al., 1999; Anthes et al., 2003; Baguette & Schtickzelle, 2003; Kon-viþka et al., 2003; Schtickzelle & Baguette, 2004). The focus on declining species, however, results in neglect of butterflies that have evaded decline and have even spread. Arguably, this restricts our understanding of species per-sistence in fragmented landscapes. It is little known, for instance, whether species avoid decline owing to less spe-cialised requirements and hence access to a wider range of biotopes, or whether they posses some demographic traits, such as better dispersal ability, which allows them to track habitable areas more efficiently. In the case of widespread generalists, a combination of the above condi-tions may be responsible, although a denser distribution of resources seems to correlate more commonly with suc-cesses (Sutcliffe et al., 1997; Schneider et al., 2003). The question is, what factors affect the apparent success of non-declining specialists.The Lesser Marbled Fritillary, Brenthis ino (Rottem-burg, 1775), represents an apparently successful species of humid meadows. Its distribution is stable in E urope. Although it has disappeared from one country (the Neth-erlands) and has declined in a few others, it is increasing elsewhere, e.g., in the Czech Republic, Hungary, Slovakia and Slovenia (Van Swaay & Warren, 1999). In the Czech Republic, the increase is accompanied by colonisation of previously unoccupied lowlands (Švestka, 1992; Beneš et al., 2002). The expansion has been attributed to expan-sion of its larval host plant, Filipendula ulmaria, facili-tated by abandonment of active management of riparian grasslands (Švestka, 1992). However, the basic life his-tory of the butterfly remains so little known that the hypothesis remains unsupported. In any case, the increase stands in striking contrast to severe declines of other spe-cies of humid meadows, such as Melitaea diamina (Lang, 1789),Euphydryas aurinia (Rottemburg, 1775) and Lycaena helle (Denis & Schiffermüller, 1775) (cf. Benešet al., 2002). B. ino thus provides a suitable model for studying mechanisms of persistence of specialised species in fragmented landscapes.Here, we present results of a study on adult demo-graphy of B. ino in a submontane landscape in western Bohemia, Czech Republic. It is based on a survey of the distribution of the butterfly in a section of landscape, mark-recapture study of adult demography, and records of adult behaviour and resource use. We ask, which lifeEur. J. Entomol. 102: 699–706, 2005ISSN 1210-5759Adult demography, dispersal and behaviour of Brenthis ino (Lepidoptera: Nymphalidae): how to be a successful wetland butterflyK AMIL ZIMMERMANN1, Z DENċK FRIC1,2, L ADISLAVA FILIPOVÁ3 and M ARTIN KONVIýKA1,2* 1Department of Zoology, School of Biological Sciences, University of Southern Bohemia, Branišovská 31, 370 05 ýeskéBudČjovice, Czech Republic2Department of Ecology and Conservation, Institute of Entomology, Czech Academy of Sciences, Branišovská 31, 370 05 ýeskéBudČjovice, Czech Republic3Department of Botany, Faculty of Sciences, Palacký University Olomouc, ŠlechtitelĤ 13, Olomouc, Czech Republic.Key words. Argynnini, Nymphalidae, Brenthis ino, behaviour, butterfly ecology, dispersal, habitat quality, grasslands, host plant range, metapopulationAbstract. Continent-wide loss of traditionally managed humid meadows is raising concern for associated butterfly specialists across Europe. However, not all species associated with this biotope are threatened, and the Lesser Marbled Fritillary (Brenthis ino) has even spread locally. We employed mark-recapture and transect walks to study its population structure and patterns of landscape occupancy in a hilly region of western Bohemia, Central E urope, to determine which life history or demography traits might be responsible for its success. A population studied by mark-recapture harboured more than 1000 individuals and was interconnected with other populations. This was further supported by a fit of the inverse power function to dispersal data. Observations of adult behaviour revealed a broad host plant range: at least three species of plants were used by the single population. On a landscape scale, the butterfly exhibited an aggregated distribution matching its host plants. It was associated with the distribution of characteristic plants and butterflies of semi-natural humid meadows, but its frequency exceeded those of other humid grasslands specialists. The relatively broad host range coupled with varying biotope requirements of individual host plant species results in a wide biotope range for the butterfly, explaining its persistence in fragmented Central European landscapes.699 *Corresponding author; e-mail: konva@entu.cas.czhistory and demography traits might be responsible for the apparent success of B. ino in fragmented landscapes of Central Europe. By reporting our behavioural observa-tions, we also fill a gap in the basic knowledge of life his-tory of the species.METHODSStudy speciesB. ino is a Palaearctic species distributed from W. Europe to the Ussuri region, N. China and Japan. Apart from the British Islands and extreme South, it inhabits entirety of E urope (Tolman & Lewington, 1997). Principal biotopes inC. and W. Europe include humid to wet grasslands, riverine marshes, bogs, clearings in wet forests, mountain valleys and subalpine tall-herb formations. In the Czech Republic, post-1980 distribution covers 270 grid squares, or 35% of the country (Beneš et al., 2002).Individual development is univoltine, adults are on the wing from late June until mid-August. Males use patrolling to search for females, females lay eggs singly and larvae feed solitarily. Regarding host plant use, there is contradictory information in the literature. The most frequently reported plant species are Filipendula ulmaria,Sanguisorba officinalis and Rubus spp., but some authors mention Aruncus vulgaris,Potentilla palustris, Sanguisorba minor, and a few other plant species (e.g., Hrubý, 1964; Henriksen & Kreutzer, 1982; SBN, 1987; Ebert & Renn-wald, 1991; Tolman & Lewington, 1997; Settele et al., 1999;Agnes, 2000; Sawchik et al., 2003). Hence, it seems that B. ino is associated with various Rosaceae, perhaps with a trophic range varying with geographic locality.Plant nomenclature follows Kubát (2002) and butterfly nomenclature follows Karsholt & Razowski (1996).Study area and designThe study was carried in the vicinity of Karlovy Vary, western Czech Republic (50°9´N, 13°2´E, altitude 650 m), on a hilly piedmont of the volcanic Doupovske Mts. The landscape is a fine-grained mosaic of both extensively used and improved meadows, pastures, ponds, small woodlots and shrubby forest-steppes on basaltic outcrops. Remnants of wet, extensively used meadows constitute a Czech stronghold for several declining butterflies, most notably Euphydryas aurinia (cf. Konviþka et al., 2003; Hula et al., 2004).We conducted two parallel surveys in summer 2004. One was a transect-based census of butterflies in a wider landscape, the other was a mark-release-recapture (MRR) survey of selected colonies. The transects intersected the area of the MRR survey (Fig. 1), facilitating a comparison of the observational and MRR results.Mark-recaptureThe MRR study was carried out in a system of semi-natural humid meadows, separated by ponds, shrubby hedges and woodlots. They are managed by mowing once a year, but less accessible or waterlogged parts, varying in extend from year to year, are temporarily left unmown. The meadows form an approximate crescent adjoining a large species-poor improved meadow; the total area was 28 ha. The centroids of thirty indi-vidual meadow fragments (mean area = 0.9 ha, SD = 0.64 ha), distinguishable by prominent landmarks, were used for ana-lysing mobility.The study was conducted from 21 June, when we observed the first individual, until 5 August. E ach day, weather permit-ting, one to three persons traversed each of the meadow frag-ments; the time spent within each fragment was proportional to fragment size. As many butterflies as possible were captured and marked with unique numbers; their sex, wing wear (1–4 scale, 1 being fresh and 4 being heavily worn) and position of capture were noted before releasing them at capture points. We also recorded behaviour prior to capture, distinguishing “flight”,“patrolling” (searching flight distinguished in males only), bask-ing, resting, egg-laying, mating and nectaring. For nectaring, we recorded the plant used.To estimate daily population sizes N i, we used the analytical Jolly-Seber method computed as model A in the program JOLLY (Pollock et al., 1990). The method is appropriate for open populations subject to births, deaths, immigration and emi-gration. The estimates are based on numbers of marked animals, estimates of daily residence (M i – this combines survival and the probability of staying in a population) and daily numbers of ani-mals at risk of being captured. It does not return a “total” popu-lation size, which would be meaningless for open population systems.To estimate average residence, we used the variance weighted averaging (Tabashnik, 1980), which weights the JOLLY-estimated j i s by the reciprocals of their standard errors V (M i)–(ln M V)–1. All the estimates were performed separately for sexes.We used the inverse power function (Hill et al., 1996; Kuras et al., 2003) to asses the probabilities of movements to distances beyond those covered by the MRR study. The function has fatter tail than an alternative, the negative exponential function, and hence is more appropriate for predicting rare long-distance movements (Baguette, 2003; Vandewoestijne & Baguette, 2004). For all butterflies captured more than once, the prob-ability density I of movements to distances D equals:nIPFDCI700Fig. 1. Map of the study area showing the humid meadows where the mark-recapture study was carried out, and the four transect routes dissecting the meadows. Thick lines show seg-ments occupied by Brenthis ino.The parameters C and n are estimated by plotting the logarithm of cumulative fractions of individuals moving specific or greater distances (ln I) against linearised expressions of the distances, i.e., ln I = ln C – n(lnD). We compared slopes and intercepts of the resulting linear regressions using t-tests (Zar, 1999). Transect surveyWe delimited four parallel transect routes, each consisting of twenty segments 300 m long and situated 300 m apart, thus forming a rectangular grid of 20 × 4 segments (Fig. 1). Pre-vailing biotopes along the routes were meadows and pastures (both unimproved and improved), plus small woodlots, scrub, and arable fields. We walked the routes approximately once every ten days between late May and early August, carrying out eight walks in all. The walks were limited to suitable weather, between 9:30 and 16:00 (C. European summer time), and it took two person-days to walk the entire grid. We counted all butter-flies and burnet moths (Zygaenidae) observed. We also recorded plant diversity along the lines by identifying to species all higher plants growing in an approximate 5 m strip along the route.We used the butterfly counts to calculate the frequency and abundance of B. ino relative to other butterflies. The pattern of aggregation along the transect was assessed by the Morisita index of dispersion (Krebs, 1989). Distributions of the butterfly and its host plants were compared using Mantel’s test for two distance matrices (Sorensen’s similarity), one based on presence-absence of B. ino and one based on presence-absence data of its host plants.Patterns of association with other butterflies, and with plants, were analysed using redundancy analysis, a linear ordination method that arranges samples according to their species compo-sition, constraining the ordination according to independent “environmental variables”. For butterflies, we used presence of individual species, less B. ino, as “species data”, and the presence/absence of B. ino as a categorical predictor. For plants, we again used species presence data as dependent variables and B. ino as a predictor. The computations were done in CANOCO for Windows 4.5 (Ter Braak & Smilauer, 2002), using the Monte-Carlo permutation test (999 runs) for assessing the sig-nificance of the ordinations, and considering the spatial struc-ture of the data in permutation designs.RESULTSDemographyWe marked 1662 butterflies and obtained 1446 recap-tures during the MRR study (Table 1). Capture sex ratio was male biased (F2 = 32.3, 1 d.f., p < 0.001), and males were recaptured more often than females (F2 = 35.5, 1 d.f., p < 0.001).The seasonal recruitment was markedly protandrous. The first male appeared 14 days earlier than the first female (21 June vs. 7 July) and the peak of male flight preceded that of females by about two weeks (Fig. 2). Male recruitment was characterised by a steep increase. The beginning of female flight was less abrupt, and we missed an expected tail of the female flight period in August. This was further evident from the pattern of sea-sonal increase of wing wear. When plotted against marking days, the increase was steeper in males (E= 0.62, F = 1287.8, d.f. = 1, 2052, p < 0.001) than in females (E = 0.32, F = 80.3, p < 0.001; comparison of the regression coefficients: t = 6.3, d.f. = 274, p < 0.001). In addition, daily residence rates tended to decrease with season in males (E = –0.01, F = 3.89, d.f. = 1, 36, p = 0.06), but not in females (F = 1.19, d.f. = 1, 25, p = 0.29). The estimated mean residence rates were 0.87 (males) and 0.80 (females), corresponding to mean residence times of 6.9 and 4.5 days, respectively.Estimates of adult numbers documented a large popula-tion size, with mean values from five peak days being 710 (51 SD) males and 1150 (249 SD) females, corresponding to peak densities of 25 males and 41 females per hectare. MobilityFemales moved among individual meadows more fre-quently than males (F2 = 15.1, d.f. = 1, p < 0.001) and covered greater distances between consecutive captures (Mann-Whitney U, Z = –2.74, p < 0.01) (Table 2). Fitting mobility using the inverse power function resulted, for males, in the equation, ln I = –3.72 (± 0.100)–1.58 (± 0.100) × ln D.The fit was highly significant (F = 126.9, d.f. = 1, 17, p < 0.001), with R2 = 0.85. For females, the equation was ln I = –3.60 (± 0.135) –1.51 (± 0.110) × ln D.It was again significant (F = 126.9, d.f. = 1, 17, p < 0.001), with R2 = 0.88.The two regressions did not differ (t = 0.47, d.f. = 45, p > 0.1), suggesting essentially identical dispersal kernels with the exponential parameter n (determining the width of dispersal tail) close to 1.5. The estimated probabilities701Fig. 2. JOLLY estimates of daily population sizes of Brenthis ino, based on a mark-recapture survey in 2004 in an area of wet meadows in western Bohemia. The error lines are standard errors of the estimates.of long-distance flights were 0.02 for 1 km, 0.002 for 3km, and 0.0006 (males) or 0.0009 (females) for 5 km.Given the dense distribution of the butterfly in the land-scape (see below), it is likely that local colonies were all interconnected by individual dispersal.Behaviour and host plant choicePrevailing behaviour of males observed during the MRR was patrolling (Fig. 3). They patrolled all day in search for females, interrupting this activity only to bask or feed on nectar, whereas females flew less frequently and spent more time feeding. This resulted in a significant difference between sexes (F 2= 283.5, d.f. = 5, p < 0.001;male patrolling was merged with flight for this test).Mating (n = 5) occurred low down (< 0.7 m) on herba-ceous vegetation; one mating observed from beginning to end lasted 140 min.E gg-laying was preceded by a short searching flight.Once a female landed on a host plant, she immediately descended towards the ground, located a suitable leaf near the ground, flexed her abdomen upwards and depos-ited one egg on leaf underside, close to the leaf base. Shethen worked her way upwards, basked for a few seconds and flew a few meters away, where she continued basking for further 8–15 min. Many of the basking or egg-laying activities observed during MRR (Fig. 3) were probably associated with oviposition.We observed 23 eggs being laid. The host plants used were Sanguisorba officinalis (19),Potentilla erecta (3)and Comarum palustre (1). The females did not seem to discriminate with respect to height of vegetation. After 16July, about two thirds of the meadows were mown, and we repeatedly observed females visiting the mown parts and ovipositing on regenerating Sanguisorba officinalis .We failed to observe oviposition on Filipendula ulmaria ,although the plant was common in the study area and formed dense growths along meadow edges. However,we often encountered females alighting on clumps of the plant. Because F. ulmaria is a tall, bulky herb, we can not exclude the possibility that it is used as host plant, but we failed to observe it.Fig. 4 presents nectaring records. Although the but-terfly uses a wide array of plants, over half of all observa-tions were on just three species in both sexes. The differ-ences between sexes in relative importance of nectar sources reflected a phenological shift in plant flowering times.Distribution in wider landscapeWe obtained 8717 records of 54 butterfly species from the transect survey. Means/medians per segment were 109(± 62.3 SD)/99 individuals, and 14 (± 4.0 SD)/14 species.The abundances per species were closely correlated with frequencies (Spearman’s s = 0.96, t = 24.0, d.f. = 52, p <0.001), and both measures were distinctly left-skewed (mean/median frequencies: 21 (± 19.3 SD)/11;mean/median abundance: 161 (± 218.3 SD)/20).Brenthis ino was observed in 36 segments (45% of the total), numbering 390 individuals (mean per segment: 5.0± 6.7 SD) and ranking as the thirteenth most frequent and ninth most abundant species. Most of the butterflies exceeding it in frequency and abundance are widespread in C. E urope (e.g. Aglais urticae ,Aphantopus hyperan-thus ,Maniola jurtina ,Coenonympha pamphilus ,Thyme-licus lineola ). The only biotope specialists exceeding B.ino in both abundance and frequency were Polyommatus amandus and Erebia medusa . Of course, the abundance ranking could have changed by extending the walks towards early May/late August, but the changes would unlikely affect the overall pattern radically. In particular,the period covered by the observations included flight periods of all wet meadows specialists.Of the 390 observations of B. ino , 122 (32%) originated from the MRR area. Considering that there were about 1900 butterflies during the peak flight time we estimated702Fig. 3. Types of behaviour prior to capture observed during the mark-recapture study of Brenthis ino .the occurrence of some 6000 butterflies in the landscape covered by the transect.The distribution of B. ino along the transect was highly aggregated (Morisita index of dispersion = 0.06). Despite this, the longest distance separating two occupied seg-ments was 1080 m, which was within a range easily crossed by dispersers. Its host plants were present in either 42 (including Filipendula ulmaria ) or 36(excluding it) segments, and the distributions of the but-terfly matched that of the plants (Mantel test, p = 0.94).The longest distances separating segments with host plants were as little as 300 m, suggesting that dispersing females should easily be able to locate host plant biotopes.Both ordination of plants (Fig. 5a) and butterflies (Fig.5b) corroborated a close association of B. ino with char-acteristic species of humid meadows. The positions of the ten most closely associated butterflies (according to spe-cies scores on the first ordination axis) on a rank-frequency plot (Fig. 6) shows that B. ino was the third most frequent associated species, surpassing, both in fre-quency and in abundance, all other characteristic wet meadow specialists.DISCUSSIONIn a submontane landscape of the western Czech Republic, Brenthis ino occurs in high densities, its abun-dance and frequency of occurrence exceed those of other characteristic butterflies of humid meadows. It exhibits a wide trophic range, its host plants are ubiquitous in the landscape. Flight distances realistically crossed by dis-persing individuals exceed the distances separating areas without host plants. The wide distribution is further facili-703Fig. 4. Use of nectar plants by males and females of Brenthis ino, recorded during mark-recapture survey.Fig. 5. Ordination diagrams (redundancy analysis) showing association of Brenthis ino with plants and butterflies recorded along transect routes. The ordinations are based on presence-absence data, with presence/absence of B. ino as an independent categorical predictor. Only species with the highest fit in the models are shown for clarity. (a) Plants: eigenvalues of the first and second axes: 0.078 and 0.037; Monte-Carlo permutation test of significance of the first axis: F = 5.68, p < 0.01; (b) But-terflies: Eigenvalues: 0.056 and 0.143; test: F = 4.59, p < 0.01.tated by such life history traits as laying eggs singly and using patrolling tactic for mate location.Our observations added two species, Potentilla erecta and Comarum palustre , to already known trophic range,further supporting the notion that B. ino feeds on a broad range of Rosaceae hosts. The locally used plants vary in biotope requirements (Kubát, 2002). Sanguisorba officinalis occurs in mown meadows with intermediate levels of nitrogen and humidity; Potentilla erecta prefers drier and nitrogen-poor mountain grasslands; and Comarum palustre is a characteristic species of peat bogs.We failed to observe oviposition on Filipendula ulmaria ,which grows at abandoned nitrogen-rich sites. Some of the plants are locally abundant. This was not the case with rare and endangered Comarum , but both Sanguisorba and Potentilla were common in the study landscape and Fili-pendula locally dominated the vegetation. Further study of potential preference for individual hosts is urgently needed.The effect of broad host plant range becomes apparent when B. ino is compared with co-occurring but declining species. At the study site, Melitaea diamina feeds solely on Valeriana dioica , a relatively rare plant (twelve tran-sect segments). Similarly, Euphydryas aurinia feeds on Succisa pratensis (nine segments), and selects plants growing in low sward and relatively low humidity (Kon-vi þka et al., 2003).The conjecture that broad trophic range and wide distri-bution of host plants translates into high local densities and wide landscape-scale distribution of a butterfly is far from trivial, because such clear-cut relationships are only rarely demonstrated. Our population used at least three (and possibly four) plants within one site. In contrast,many butterflies with seemingly broad host ranges differ in host use among populations (e.g., Kuussaari et al.,2000; Wahlberg, 2001). Syntopic feeding on a wider range of plants is typical for generalists belonging to such groups as pierids, but seems to be rare among Argynnini fritillaries. It is noteworthy that another fritillary that has been recently expanding in the Czech Republic, Proclos-siana eunomia , follows local increases of its host plant,Bistorta major , caused by abandonment of mountain meadows (Pavli þko, 1996). Similarly, the only “large”fritillary still relatively common in the country, Argynnis aglaja , utilises a broader host range than its declining congenerics (Fric et al., 2005).Dispersing individuals of B. ino easily locate habitable patches. There is an argument strongly suggesting that a dense distribution of resources is more important than innate mobility. In an inverse power function, the expo-nential parameter n determines the length of frequency-distribution tail: the lower its value, the higher the dis-tances that are crossed (Hovestadt et al., 2001). Consid-erably lower values than those detected for B. ino were reported, e.g., for the bog fritillary Boloria aquilonaris in Belgium, where it is threatened (Baguette, 2003), or for a widespread grassland satyrid Melanargia galathea (Van-dewoestinje & Baguette, 2004). Values ranging from 1 to 2 (close to 1.5 in B. ino ) seem to be more frequent. How-ever, they occur again both in declining and widespread species. They were found, among others, for Euphydryas maturna , the most severely threatened butterfly in the Czech Republic (Konvi þka et al., 2005). Within our MRR area, such values applied for the declining Euphydryas aurinia and Melitaea diamina , as well as for the still common M. athalia (unpubl. data). High propensity to move cannot generate a wide distribution under scarcity or absence of resources. Instead, species of intermediate mobility tend to suffer most severely if affected by habitat loss (Thomas, 2000).The efficient dispersal of B. ino may be influenced by some subtler traits. Norberg et al. (2002) demonstrated that B. ino crossed shaded areas more readily than four other grassland species tested in a simultaneous outdoor-cage experiment, perhaps owing to its larger size and hence more efficient thermoregulation (cf. Heinrich,1986).Another factor possibly contributing to the success of B. ino is its low selectivity for habitat architecture. Its males patrol, and hence can locate females over wide tracks of uniformly mown meadows. This contrasts with at least two other wet meadows butterflies, Lycaena helle and Euphydryas aurinia ,which require meadows dis-sected by wind-shielded structures such as hedges, on which their males establish perches (Fisher et al., 1999;and unpublished observation). Furthermore, females of B.ino lay eggs singly, spreading the risks faced by progeny of individual females. Under such a strategy, it is unlikely that too exacting preferences for plants growing in par-ticular conditions would have evolved. This is supported by our observation that they oviposited on both unmown and freshly mown plant individuals.The number of eggs produced per female depends on female longevity. Argynnini relatives of B. ino are typical704Fig. 6. Rank-frequency plot for butterflies observed during the transect census in piedmont of Doupovske Mts, western Bohemia, showing the frequency of Brenthis ino (filled circle)relative to associated species. Ten species that exhibited closest association with B. ino in a redundancy analysis are shown as filled diamonds; characteristic species of humid meadows are underlined.income breeders, whose females derive resources for egg production from adult diet (Boggs & Ross, 1993; Boggs, 1997; O’Brien et al., 2004). In our study, the residence values for females were shorter than for males, but the estimates were biased due to the missing tail of female emergence. On the last marking day, average female wing wear was just 2.4 (n= 14), contrasting with 4.0 in males (n = 3), suggesting that many females would live for sev-eral more days. This and the maximum values for observed residency document that adults of B. ino are relatively long-lived, consistent with their income-breeding strategy (Boggs, 1997; Kopper et al., 2001).To summarise, the remarkably broad host plant range combined with a wide distribution of the host plants seems to be the primary factor facilitating the persistence of Brenthis ino in western Bohemia, and elsewhere in E urope (cf. Saarinen et al., 2003; Sawchik et al., 2003). Demographic and life history traits likely contribute to the success, but probably play a rather minor role. Avail-ability of extensively used hay meadows with infre-quently mown edges is crucial for further persistence of the species.ACKNOWLEDGEMENTS. We thank to M. Bartoš, V. ýihák-ová, V. Hula, M. Klímová and M. Maradová for help in field. The study was supported by the Czech Academy of Sciences (B6007306/2003) and by the Czech Ministry of E ducation (6007665801).REFERENCESA GNES G. 2000: Schmetterlingszönosen des Feuchtgrünlandes in der Deutsch-Belgischen Hocheifel und U ntersuchungen zur Eignung von Indikatorarten für die Differenzierung vernäßter Standorte. Diploma thesis, Institut für Evolutionsbiologie und Ökologie, Universität Bonn, Bonn, 83 pp. On-line: http://www.gregoragnes.de/Diplom/Diplomarbeit.pdfA NTHES N., F ARTMANN T., H ERMANN G. & K AULE G. 2003: Com-bining larval habitat quality and metapopulation structure –the key for successful management of pre-alpine Euphydryas aurinia colonies. J. Insect Conserv.7: 175–185.B AGUETTE M. 2003: Long distance dispersal and landscape occu-pancy in a metapopulation of the cranberry fritillary butterfly. Ecography26: 153–160.B AGUETTE M. & S CHTICKZELLE N. 2003: Local population dynamics are important to the conservation of metapopula-tions in highly fragmented landscapes. J. Appl. Ecol. 40: 404–412.B ENEŠ J., K ONVIýKA M., D VOěÁK J., F RIC Z., H AVELDA Z., P AV-LIýKO A., V RABEC V. & W EIDENHOFFER Z. 2002: [Butterflies ofthe Czech Republic: Distribution and Conservation] I., II. SOM, Prague, 857 pp. (in Czech, English summaries).B OGGS C.L. 1997: Reproductive allocation from reserves and income in butterfly species with differing adult diets. Ecology 78: 181–191.B OGGS C.L. & R OSS C.L. 1993: The effect of adult food limita-tion on life history traits in Speyeria mormonia (Lepidoptera, Nymphalidae).Ecology74: 433–441.E BERT G. & R ENNWALD E. 1991: Die Schmetterlinge Baden-Württembergs,Band 1: Tagfalter 1. E ugen Ulmer, Hohen-heim, 552 pp.F ISHER K., B EINLICH B. & P LACHTER H. 1999: Population struc-ture, mobility and habitat preferences of the violet copper Lycaena helle(Lepidoptera, Lycaenidae) in Western Ger-many – Implications for conservation. J. Insect Conserv.3: 43–52.F RIC Z., K LÍMOVÁ M., H ULA V. & K ONVIýKA M. 2005: Caterpil-lars of Argynnis aglaja(Linnaeus, 1758) (Lepidoptera, Nym-phalidae) feeding on Bistorta major. Atalanta36: 119–121.H EINRICH B. 1986: Comparative thermoregulation of 4 montane butterflies of different mass. Physiol. Zool.59: 616–626.H ENRIKSEN H.J. & K REUTZER I. 1982: The Butterflies of Scandi-navia in Nature. Skandinavisk, Bogforlag A/S, Odense, 215 pp.H ILL J.K., T HOMAS C.D. & L EWIS O.T. 1996: Effects of habitat patch size and isolation on dispersal by Hesperia comma but-terflies: Implications for metapopulation structure. J. Anim. Ecol.65: 725–735.H OVESTADT T., M ESSNER S. & P OETHKE H. J. 2001: Evolution of reduced dispersal mortality and “fat-tailed” dispersal kernels in autocorrelated landscapes. Proc. R. Soc. London (B)268: 385–391.H RUBÝ K. 1964: Prodromus Lepidopterorum Slovaciae. SAV, Bratislava, 962 pp.H ULA V., K ONVIýKA M., P AVLIýKO A. & F RIC Z. 2004: Marsh Fritillary (Euphydryas aurinia) in the Czech Republic: Moni-toring, metapopulation structure, and conservation of the endangered butterfly. Entomol. Fenn. 15: 231–241.K ARSHOLT O. & R AZOWSKI J. 1996: The Lepidoptera of Europe, a Distributional Checklist. Apollo Books, Stenstrup, 380 pp.K ONVIýKA M., H ULA V. & F RIC Z. 2003: Habitat of pre-hibernating larvae of the endangered butterfly E uphydryas aurinia(Lepidoptera: Nymphalidae): What can be learned from vegetation composition and architecture? Eur. J. Ento-mol.100: 313–322.K ONVIýKA M., ýÍžEK O., F ILIPOVÁ L., F RIC Z., B ENEŠ J., K RUPKA M., ZÁMEýNÍK J. & D OýKALOVÁ Z. 2005: For whom the bells toll: Demography of the last population of the butterfly E uphydryas maturna in the Czech Republic. Biologia (in press).K OPPER B.J., M ARGOLIES D.C. & C HARLTON R.E. 2001: Life his-tory notes on the regal fritillary, Speyeria idalia (Drury) (Lepidoptera: Nymphalidae), in Kansas tallgrass prairie. J. Kans. Entomol. Soc.74: 172–177.K REBS C.J. 1989: Ecological Methodology. Harper & Collins, New York, 654 pp.K UBÁT K. (ed.) 2002: Klíþ ke kvČtenČýeské republiky[Key to Flora of the Czech Republic]. Academia, Praha, 927 pp. (in Czech).K URAS T., B ENEŠ J., F RIC Z. & K ONVIýKA M. 2003: Dispersal pat-terns of endemic alpine butterflies with contrasting population structures: Erebia epiphron and E. sudetica. Popul. Ecol.45: 115–123.K UUSSAARI M., S INGER M. & H ANSKI I. 2000: Local specializa-tion and landscape-level influence on host use in an herbivo-rous insect. Ecology81: 2177–2187.M AES D. & V AN D YCK H. 2001: Butterfly diversity loss in Flan-ders (north Belgium): E urope’s worst case scenario? Biol. Conserv.99: 263–276.N EVE G., B ARASCUD B., H UGHES R., A UBERT J., D ESCIMON H., L EBRUN P. & B AGUETTE M. 1996: Dispersal, colonization power and metapopulation structure in the vulnerable but-terfly Proclossiana eunomia (Lepidoptera: Nymphalidae). J. Appl. Ecol.33: 14–22N ORBERG U., E NFJÄLL K. & L EIMAR O. 2002: Habitat exploration in butterflies – an outdoor cage experiment. Evol. Ecol.16: 1–14.O’B RIEN D.M., B OGGS C.L. & F OGEL M.L. 2004: Making eggs from nectar: the role of life history and dietary carbon turn-705。
