水域生態(tài)學：Ch3 群落生態(tài)學Community Ecology

1、DefinitionInterspecific interactionsCommunity structureCommunity successionCh. 3 群落生態(tài)學Community Ecology1. 群落定義在同一時間、同一地方一起出現(xiàn)的所有物種種群的集合。An assemblage of species populations that occur together in the same place at the same time. 一個生物群落是生活在同一環(huán)境中、相互支持、相互之間有直接或間接作用的植物和動物的自然集合。A biotic community is a nat

2、urally occurring assemblage of plants and animals living in the same environment, mutually sustaining and interacting directly or indirectly with one another.群落定義植物、動物、微生物通過攝食關系和別的作用相互關聯(lián)，形成一個綜合整體，稱之為生物群落。these plants, animals, and microbes are linked to one another by their feeding relationships and

3、 other interactions, forming a complex whole referred to as a biological community群落的不同觀點整（機）體論holistic concept(Clements 1916,1936) 認為群落是一個超生物體(superorganism)，只有將其作為一個完整的實體考慮時，才能正確地評價其功能和組織。只有依據(jù)各個物種對整個系統(tǒng)動態(tài)所作的貢獻，才能了解每個物種。個體論 individualistic concept (Gleason 1926,1939)認為群落結構和功能僅僅表達各物種的相互作用，并未反映任何超越物種水

4、平之上的組織。群落的不同觀點整體觀預言群落是封閉的(closed) ：物種分布是同時發(fā)生的(coincident)群落間邊界(群落交錯區(qū) ecotones)是明確的distinct個體觀預言群落是開放的open :物種分布是不相關的(independent)群落間邊界是模糊的(diffuse) 群落的不同觀點中間的觀點或混合的觀點接受個體論的觀點：大多數(shù)的相互作用是敵對的，群落是物種偶然的集合； 接受整體論的觀點：群落的一些特性來自于物種間的相互作用，由于協(xié)同進化而加強。2. 種間關系Interspecific interactions 積極關系Positive interactions互利共

5、生(Mutualism)偏利共棲 (Commensalism)促進作用 (Facilitation): Enhancement of a population of one species by the activities of another, particularly during early succession. 種間關系消極關系Negative interactions競爭 (Competition )捕食 (Predation)寄生 (Parasitism)偏害作用 (Amensalism): A symbiotic relationship between organisms

6、in which one species is harmed or inhibited and the other species is unaffected. 抑制作用 (Inhibition): the suppression of a colonizing population by another that already established, especially during successional sequences. 2.1 互利共生 (Mutualism)Mutualism: 雙方互利的共生合作關系 (The type of symbiosis in which bot

7、h species profit from the association). 專性互利共生obligate when the participating species are fully dependent on one another 兼性互利共生facultative when either partner can live alone if necessary Anthozoan(珊瑚蟲)zooxanthella (蟲黃藻)；cleaner fishcustomer fish Mutualism two species provide resources or servicesto

8、each other enhances fitness of both speciesMutualism. The greenish-brown specks in these coral polyps are zooxanthellae, algae that live symbiotically within the corals tissue and supply the coral with carbon and nitrogen compounds. In return, the coral provides its zooxanthellae with nitrogen in th

9、e form of ammonia.Mutualism: cleaner wrasse (隆頭魚) and sea bass (鮨)Mutualism: hummingbird and flower2.2 偏利共棲 (Commensalism)Commensalism：一方得利，另一方無利也無害的共生關系 (The form of symbiosis in which one species benefit from the association, while the other is neither harmed nor benefited).Commensalism one specie

10、s receives a benefit from another species enhances fitness of one species; no effect on fitness of the other speciesCommensalism. Fishes, such as this individual of Amphiprion periderion (雙鋸魚) in Guam, often form symbiotic associations with sea anemones (?？?, gaining protection by remaining among th

11、eir tentacles and gleaning scraps from their food2.3 競爭Competition 資源利用性競爭 (Exploitation competition ): 一個物種的競爭行為顯著減低其他物種對共享資源的可得性相互干涉性競爭(Interference competition): competition in which individuals actively interfere with one another access to resources. Individuals interact directly, most obviously

12、, in the case of some animal species, by fightingby producing toxins (e.g. plant allelopathy : direct inhibition of one species by another using noxious or toxic chemical.) 競爭方式消費競爭(Consumptive competition), based on the utilization of some renewable resources搶先競爭(Preemptive competition), based on t

13、he occupation of open space蔓遮競爭(Overgrowth competition), which occurs when one individual grows upon or over another, thereby depriving the second of light, nutrient-laden water, or some other resource競爭方式他感作用(克生作用Chemical competition /allelopathy), by production of a toxin that acts at a distance a

14、fter diffusing through the environment。某些沉水植物分泌物對藻類有抑制作用領域競爭Territory competition, the defense of space遭遇競爭Encounter competition, involving transient interaction over a resource that may result in physical harm, loss of time or energy, or theft of foodStrongest competition is between similar species

15、: ecological niche ！生態(tài)位 (Niche)定義： 一個物種在群落中的生態(tài)角色 (The ecological role of a species in the community)生物或物種能維持存在的各種環(huán)境條件和資源，是一個多維空間 (the many ranges of conditions and resource qualities within which the organism or species can persist, often conceived as a multidimensional space。 Hutchinsonian niche, h

16、ypervolume niche)生境(空間)生態(tài)位habitat niche功能生態(tài)位functional niche (營養(yǎng)生態(tài)位 trophic niche)生態(tài)位基礎生態(tài)位 (Fundamental niche ) ：生物在環(huán)境中沒有其他物種作用的情況下所處的生存條件范圍。The range of condition under which an organism could survive if it were only one in the environment 實際生態(tài)位 (Realized niche)：生物實際占有的生態(tài)位，它不同于基礎生態(tài)位是因為有其他物種的存在。The

17、actual niche occupied by an organism; it differs from the fundamental niche because of the presence of other species. 生態(tài)位生態(tài)位寬度(Niche breadth): 個體、種群和物種所利用資源的多樣性和耐受條件的范圍。The variety of resources utilized and range of conditions tolerated by an individual, population, or species.生態(tài)位重疊(Niche overlap):

18、兩個或更多個物種共享生態(tài)位空間；資源需求和生態(tài)條件耐受限的相似性。The sharing of niche space by two or more species; similarity of resource requirements and tolerance of ecological conditions.生態(tài)位生態(tài)位補充(Niche complementarity): 物種在一種資源上利用上有很大重疊，必然在其它資源利用上有相當大的分化。A situation in which species that overlap extensively in their use of one

19、 resource differ substantially in their use of another resource. 生態(tài)位先占(Niche preemption): 先占的物種在一個地區(qū)相繼獲得一部分可得資源，那么留給后來移殖的物種的資源就較少。A model in which species successively procure a proportion of the available resources in an area, leaving less for the next colonizing species. 生態(tài)位生態(tài)位壓縮(niche compression

20、 /contract)：競爭導致生境壓縮，而不引起食物類型和所利用資源的改變。競爭釋放(competitive release)：由于種間競爭減弱而引起生態(tài)位的擴展生態(tài)位移動(niche shift)：兩個或多個物種由于減弱了種間競爭而發(fā)生的行為變化和取食格局變化(高斯)競爭排斥原理 (Causes competitive exclusion principle)：占有相同生態(tài)位的兩個物種不能永遠共存 (two species with identical niches cannot coexist infinitely).P.aurelia雙核草履蟲，P.caudatum尾草履蟲避免競爭 A

21、voiding competition自然選擇傾向于通過生態(tài)位分化 (niche differentiation) 避免競爭時間和空間上分化食性上分化the absence of competition may be the “ghost of competition past”reduced niche overlapAnolis lizards: resource (or niche) partitioning: different perches 避免競爭 Avoiding competition性狀替換 (character displacement)：當現(xiàn)實生態(tài)位在競爭影響下被壓縮，

22、隨著對新的資源的適應，形態(tài)性狀跟著發(fā)生改變。如兩種達爾文雀Gespiza fortis 和G. fuliginosa 單獨出現(xiàn)時，鳥喙大小相似；同時出現(xiàn)時，G. fuliginosa 的喙明顯比G. fortis窄。 競爭和移居的權衡Competition-colonization tradeoffsimportance of disturbance, and of variability (heterogeneity) in the environmentcompetition-colonization tradeoff: some species are good at colonizin

23、g (fecund, long-dispersing seeds), while others are good at competing (preempting resources)生態(tài)位分化理論在漁業(yè)上的應用混養(yǎng)(Polyculture): 利用不同魚類棲息空間和食物需求的差異，充分利用水體資源，提高產量移植(Transplantation): 利用空的生態(tài)位(vacant niche).生態(tài)等值 (Ecological equivalents) 在不同地理區(qū)域占據(jù)相同生態(tài)位的物種。Species that occupy the same niche in different geogra

24、phical regions. Lotka-Volterra 競爭模型擴展logistic方程: 競爭系數(shù)competition coefficient 與生態(tài)位重疊有關niche overlapThe orientation of isoclines for zero population growth and the outcome of competition according to the Lotka-Volterra competition modelLotka-Volterra 競爭模型: 結論假如物種A的對角線在物種B的對角線之上，物種A就贏環(huán)境容納量大，競爭有利(r-選擇的物

25、種在這種情況下贏不了)只有在種間競爭小于種內競爭時，才能出現(xiàn)共存: 12 21 R異養(yǎng)演替(Heterotrophic succession): PR演替類型原生演替(Primary succession): 在新形成的基質上，如熔巖、新裸露的巖地、沖刷沉積物和冰磧等，無有機質存在，尚未被任何生物占據(jù)之處開始的演替次生演替(Secondary succession): 在植被被人類、動物或火、風、暴雨等擾動過的地方開始的演替。生態(tài)演替機制 驅動生態(tài)演替的機制包括：促進 (Facilitation )：體現(xiàn)Clements的發(fā)育序列觀點，每一物種為下一物種鋪平道路。抑制 (Inhibition)

26、： 一個物種的存在抑制另一物種是常見的現(xiàn)象，當抑制起作用時，演替朝著只有通過定居個體死亡和替代，導致不可避免地向長壽物種轉變。耐受(tolerance)：通過耐受，一個物種能侵入新的生境，不管是否存在其他物種而定居下來。在此機制下:早期階段是壽命短暫、競爭力弱的競爭者占優(yōu)勢；優(yōu)秀的競爭者構成頂極群落的物種，但增長緩慢后來才表現(xiàn)出優(yōu)勢來。合條件的任何種類都有機會生存干擾造成較大的空白地帶只有某些先鋒種類能夠生存先到的種類改變環(huán)境使后續(xù)種類易于移入先到的種類改變環(huán)境對后續(xù)種類移入沒有影響或影響不大先到的種類改變的環(huán)境不適合后續(xù)種類的移入耐受模型抑制模型通過競爭后續(xù)種類淘汰先到種類只要先到種類不受損

27、害且繼續(xù)繁殖，將排斥或抑制其它種類若外界干擾使原有種類受損或死亡，則由耐受性更強的種類去替代上述過程反復進行，直至現(xiàn)存種類不再促進其它種類的移入和生長，或再沒有其它種類能夠移入和生長演替機理的三種模型（仿Connell and Slatyer 1977）促進模型Eventually resident species are ones able to tolerate environmental change by earlier species and no other species can tolerate conditionsSpace available for colonizatio

28、nOnly early successional species can establishAny species able to survive as adults establishFirst species to establish modify environmentEnvironment less suitable for early species but more suitable for late successional species; early successional species die outEnvironment less suitable for early

29、 species but neither less nor more suitable for late successional speciesEnvironment less suitable for establishment by all speciesEventually resident species are ones that do not change environment in a way to favor other speciesResident species inhibit establishment of all other species; persist until disturbedDisturbance destroys climax stageFacilitationToleranceInhibition“Climax”“Climax”“Climax”Succession beginsAlternative successional mechanisms (data from Connell and Slatyer 1977)頂極群落和演替中群落的比較特征(尚玉昌、蔡曉明，1992)群落特征演替中群落頂極群落群落能量學1.總生產量/群落呼吸(P/R)112.總生產量/生物