Урбанизация «диких» видов птиц в контексте эволюции урболандшафта



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Птицы
Амадина зебровая (Taeniopygia guttata) - 40

Бормотушка северная (Hippolais caligata) – 69,

Вертишейка (Yunx torquilla) – 26,

Воробей полевой (Passer montanus) – 32, 41, 100, 101

Воробей домовый (Passer domesticus) – 33, 41, 101

Воробей саксаульный (Passer ammodendri) -33

Ворон (Corvus corax) – 25, 26, 101

Ворона большеклювая (Corvus macrorhynchos) - 102

Ворона серая (Corvus cornix) – 48, 58, 73, 86, 96, 101

Ворона чёрная восточная (Corvus corone orientalis) – 66

Вьюрок канареечный (Serinus serinus) - 100

Вяхирь (Columba palumbus) – 9, 60, 98

Галка (Corvus monedula) – 25, 26, 58

Горихвостка обыкновенная (Phoenicurus phoenicurus) – 26, 87

Горлица кольчатая (Streptopelia decaocto) – 74, 90, 99

Городская ласточка (Delichon urbica) – 25

Грач (Corvus frugilegus) – 11, 25

Дрозд-белобровик (Turdus iliacus) – 47, 88

Дрозд-деряба (Turdus viscivorus) – 88-89

Дрозд-рябинник (Turdus pilaris) – 26, 47, 61, 87, 88-89, 98, 99

Дрозд чёрный (Turdus merula) – 49, 60, 87, 88, 90, 98, 99, 102-103

Дрозд певчий (Turdus philomelos) – 85, 87, 88-89

Дубонос обыкновенный (Coccothraustes coccothraustes) - 69

Дятел большой пёстрый (Dendrocopos major) – 60, 76, 85, 86, 98

Дятел зелёный (Picus viridis) – 69,

Дятел золотой (Colaptes auratus) – 40

Дятел малый пестрый (Dendrocopos minor) – 26, 60

Дятел седой (Picus.canus) – 69

Дятел сирийский (Dendrocopos syriacus) - 98

Желна, чёрный дятел (Dryocopus martius) - 80

Зарянка (Erithacus rubecula) - 60

Зеленушка (Carduelis chloris) – 26, 55, 99

Зяблик (Fringilla coelebs) – 60, 87

Камышевка дроздовидная (Acrocephalus arundinaceus) – 69,

Камышевка тростниковая (Acrocephalus scirpaceus) – 69

Клёст-еловик (Loxia curvirostra) - 98

Конёк лесной (Anthus trivialis) - 87

Королек желтоголовый (Regulus regulus) - 26

Крапивник (Troglodytes troglodytes) – 26

Крачка речная (Sterna hirundo) – 89, 98

Кряква (Anas platyrhynchos) – 58-59, 88, 96, 101, 102

Кукушка (Cuculus canorus) – 26

Лазоревка голубая (Parus caeruleus) - 60

Мухоловка серая (Muscicapa striata) – 26

Мухоловка-пеструшка (Ficedula hypoleuca) – 52, 54-56, 60, 79, 87, 96

Нырок красноголовый (Aythya ferina) - 98

Овсянка-дубровник (Emberiza aureola) - 61

Овсянка обыкновенная (Emberiza citrinella) - 87

Овсянка садовая (Emberiza hortulana) – 69

Овсянка шалфейная (Amphispiza belli) – 79

Пеночка-весничка (Phylloscopus trochilius) - 79

Пеночка зелёная (Phylloscopus trochyloides) – 61, 69, 78

Пеночка-теньковка (Phylloscopus collybita) – 26, 98

Перепел японский (Coturnix japonica) - 86

Пищуха (Certhia familiaris) – 26

Поганка большая, чомга (Podiceps cristatus) – 98

Поганка черношейная (Podicceps nigricollis) - 98

Поползень (Sitta europaea) - 93

Сверчок соловьиный (Locustella luscinioides) - 69

Сизый голубь, сизарь (Columba livia) – 31, 46, 50, 54, 58, 76, 79, 90-91, 94, 98, 101

Синехвостка (Tarsiger cyanurus) - 61

Синица большая (Parus major) – 34, 46, 48, 51-53, 60, 76, 82, 87, 88, 93

Синица-московка (Parus ater) – 26, 93, 98

Синица-пухляк, буроголовая гаичка (Parus montanus) – 26, 93, 98

Cиница хохлатая, гренадёрка (Parus cristatus) - 98

Сипуха (Tyto alba) – 25

Скворец (Sturnus vulgaris) – 25, 96, 100

Славка садовая (Sylvia borin) – 87

Славка серая (Sylvia communis) -87

Славка ястребиная (Sylvia curruca) – 26, 69, 87

Сова ушастая (Asio otus) -60, 85

Сойка (Garrulus glandarius) – 26

Сорока (Pica pica) – 41, 86, 90, 99

Трупиал краснокрылый (Agelaius phoeniceus) - 78

Трясогузка белая (Motacilla alba) – 26

Трясогузка желтоголовая (Motacilla citreola) - 69

Чечевица обыкновенная (Carpodacus erythrinus) – 61

Чайка озёрная (Larus ridibundus) - 98

Чайки Larus spp. – 86

Чернеть хохлатая (Aythya fuligula) - 98

Чиж (Spinus spinus) – 26, 98

Хохлатый жаворонок Galerida cristata – 25

Юнко (Junco hyemalis) - 38

Ястреб-тетеревятник Accipiter gentilis – 11, 25, 61


Млекопитающие
Белка обыкновенная (Sciurus vulgaris) - 86

Долгопят бирманский (Tarsius bancanus) – 42

Крыса серая (Rattus norvegicus) - 44

Луговая собачка чернохвостая (Cynomys ludovicianus) - 40

Мышь домовая (Mus domesticus) – 44

Мышь малая лесная (Sylvaemus uralensis) – 91-92, 93

Мышь полевая (Apodemus agrarius)90

Полёвка рыжая (Gletrionomys glareolus) – 38, 91-92, 93

Полёвка красная (Gletrionomys rutilis) – 75

Полёвка тёмная (Microtus agrestis) - 92



Человек разумный (Homo sapiens) – 57, 71, 89
English Summaries
Chapter I. Urbanization processes in wild bird & mammal species: transformation of metapopulation system or adatations on individual level?
The present research deals with urbanization of wild bird and mammal species. Forms and mechanisms of population steadiness in the urban landscape have been examined. The urbanization process turned out to be a directed change of the population system forming de novo in the urbolandscape leading to a sustainable organization peculiar for the particular environment. The population organization of different types in urbolandscape is found to provide its stability under conditions of directed and fast changes accompanied with instability and heterogenous structure of habitats. It is shown that the same type of population organization meets the corresponding demands among different species settling in the urban environment. Its features are "openness" and "flowage" of the groups, far order of settlement levels and other units of population system, constant movements of the individuals between the groups as a respond to the signals of urboenvironment significant changes. The "urban" variant of the population system organization turns out to be opposite to that of the same species in the non-urban habitats. After formation of the urban types by the species and successful developing of the town, the urban population becomes separated from the maternal local population and begins to exist independently in the urban landscape. The variety of adaptation aberrations in ecology, behavior, and mode of life of urban birds is the population system stability function in the urban landscape and is not a result of individual selection. It is shown that the urbanization process of the species goes firstly on the population level being the system structure transformation developed by the species towards the most stable state in the town (city) territory. Only after the appearance of stable urban population, the urban individuals show the rapid growth of different changes in ecology, behavior, mode of life that was traditionally described by naturalists as species adaptation to the city conditions. The key features of urban population stability/instability are described. Their application to closely related species allows us to distinguish potential urbanists from unstable and vulnerable species that could be soon pushed out of the city. The application of corresponding criteria to the urban populations of such species constituting one guild allows us to predict if their developing in the given town would be successful or unsuccessful. The latter is very important since close species are, as a rule, ecologically indistinguishable in the urbanized landscapes. So one can not predict successful/unsuccessful urbanization taking into account the differences in the range of habitats, breeding success, and other external features.
Chapter II. Steadiness mechanisms in urbanized bird populations: serfing on waves of environmental changes
The present research deals with urbanization of wild bird and mammal species. Forms and mechanisms of population steadiness in the urban landscape have been examined. The urbanization process turned to be the directed change of population system forming de novo in the urbolandscape to the side of stable organization variants special for the given environment. The population organization of different types in urbolandscape is found to provide its stability in the conditions of directed and fast changes together with instability and heterogenic structure of habitats.

It is shown that among different species settling in the urban environment one and the same type of population organization meets the corresponding demands. Its features are “openness” and “flowage” of the groups, far order of settlement levels and other units of population system, constant movements of the individuals between the groups as a respond to the signals of urbo-environment significant changes. The “urban” variant of population system organization turns to be completely opposite to the population structure of the same species in the non-urban habitats. After formation of the urban types by the species and successful developing of the town the urban population separates from the maternal local population and begins to exist independently in the urban landscape.

The variety of adaptation aberrations in ecology, behavior and mode of life of urban birds is the population system stability function in the urban landscape and not the results of individual selection. It is shown that the urbanization process of the species goes firstly on the population level being the system structure transformation towards the most stable state in the town (city) territory developed by the species. Only after the appearance of stable urban population the urban individuals show the rapid growth of different changes in ecology, behavior, mode of life that was traditionally described by naturalists as species adaptation to the city conditions.

The key features of urban population stability/instability are described. Their application to relative species allows us to distinguish potential urbanists from instable and vulnerable species that could be soon pushed out of the city. The application of corresponding criteria to the urban populations of relative species constituting one guild allows us to predict if their developing of the given town would be successful/unsuccessful. The latter is very important since in urbanized landscapes close species are ecologically indistinguishable as a rule. So one can not predict successful/unsuccessful urbanization taking into account the differences in the range of habitats, breeding success, and other external features.


Chapter III. Ecological and evolutionary consequences of urbanization in birds and mammals
Ecological and evolutionary consequences of urban bird population steadiness developing during urbanization of «wild» species» were investigated. It is shown that population system steadiness determine the diversity of adaptive «aberrations» in ecology, behaviour, life-history this species which always observe in their urbanized populations.

The diversity of «aberration» (and polymorphism by adaptive characters too) increase only in periods in which population is steady and their sustainability is in progress. The increasing of frequencies this «aberration» in urbanized birds population depends only on level the steadiness of species population in this city and not on ecological and behavioural plasticity of individuals or their possibilities of adaptation to urban environment.

For example, all of this «aberrations» disappeared in period when urban population decrease through just this period they may to stopped population decreasing. Were revealed the positive feedback between growth of urbanized population steadiness in same city and accelerating urbanization process for this species in next period. On account to impact this feedback on population level, urbanization proceed rapidly and directly, it is an autocatalytic process for all «wild» species which ensure the sustainability their population system in settling city. Some criteria of urban population steadiness were established (and opposite symptoms of population unsteady). There are:

1) absence «the limit of growth» urbanized birds populations can to exponentially increase their number in limited space the «archipelago» of urban habitats; 2) birds from sustainable urban populations didn’t leave the former nesting area after transformation or destroying their typical habitats. They try to stay on the breeding in this range and began to nest in non-typical habitats, build the nest with deviate methods and house it in non-typical places.

3) number dynamic of steady populations are independent from changes of some environment factors which are essential from this species (temperature, food abundance and so on). This dependence appear at once when it turned out that population system is unsteady.

4) Birds in sustainable urbanized population will divide into two or more alternative life-strategies on the basis of population division on cohorts «moreover-settled» and «moreover-nomadic» individuals.

5) The dynamical equilibrium the frequencies of alternative strategies in urbanized population keep one’s balance in relatively stable urban environment but continually changed under impact of fluctuations urban environment or direct evolution of urban landscape.

At the expense of dynamical equilibrium between some alternative strategies urbanized population are sustainable in conditions of unstable and changing environment. permanently o

So, the urbanization of «wildlife» birds species is a classical case of elementary evolutionary appearance by N.V.Timofeeff-Ressovsky: during successful occupying the cities all of «wild» species became new type of organization in it’s population system, which can be steady in extreme level of instability, variability and heterogeneity of urban environment.

Successful urbanization process run to splitting all population this species in to parts (urban and «rural» or «native») which are not communicating vessels. We established significant distinction between urbanization possibilities of near species (between ecological similar to) which distinguished from different type of organization the population systems. Causal connections between organization type of species population systems and urbanization potential were discussed as well as ecological and evolutionary consequences of urbanization.


Chapter IV. Prognosis of bird species preservation in Moscow fauna during urban landscape development in the future (year 2000).
Chapter V. Verification our prognosis about transitions in Moscow birds fauna after 7 years ago.


1 Искусственно созданная популяция, основу которой составили свободноживущие птицы из Московского зоопарка

2 Популяция создана искусственно (выпуски на Останкинских прудах), весь период существования не имела «подпитки» из региона. Приток птиц со стороны начался только в последние годы, когда гоголь стал регулярно зимовать на Москве-реке (от Курьяновской плотины до г.Бронницы).

3 В эти годы сформировалась популяция коноплянки на Ленинских горах, см. работы К.Н.Благосклонова (1991), личн.сообщение В.Н.Калякина

4 Личн.сообщение Б.Л.Самойлова

5 Серые куропатки встречались на Люблинских и Люберецких полях орошения в 1960-1970-х гг., но в 1980-х гг. достоверно отсутствовали, вновь начав гнездиться в следующее десятилетие (Ерёмкин, 2004).

6 Формирование городской популяции затруднено из-за сокращения численности в ближних пригородах, продолжающегося сокращения площадей травянистых местообитаний в самом городе.

7 В 1962 году выводок серой утки наблюдали в Серебряном бору (тогда это была практически загородная территория). Б.Л.Самойлов, личн.сообщение.

8 И главное, увеличение межгрупповой дисперсии средних значений всех соответствующих показателей.

9Близкими считали виды, таксономически близкие (обычно одного рода), сходные между собой морфоэкологически и трофически. Практически всегда составляют одну гильдию, то есть «расходятся» по одному ресурсному измерению, а в городе вообще почти идентичны.

10 Её можно обозначит как сохраняемость вида в составе городской фауны – вероятность невымирания вида в городских местообитаниях на определённый период времени вперёд.

11 Раздел написан Г.С.Ерёмкиным и Д.М.Очаговым (ВНИИ Природа). Как отдельная публикация он была подготовлен для сборника трудов III Совещания по редким видам птиц Нечернозёмного центра России: прошедшего в г. Москве в феврале 2000 г., но по независящим от авторов обстоятельствам не была опубликована.


12 Раздел написан Г.С.Ерёмкиным


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