互利共生（如螞蟻和蚜蟲）或掠食（如瓢蟲和蚜蟲）關係為重要的生物互動關係，並且能影響生物群聚的結構與穩定生態系統。在氣候變遷持續進行下，目前已經有許多文獻探討暖化如何影響生物的互動關係。然而，有兩個重要議題仍待釐清：第一是互利共生與掠食關係經常被單獨地研究，但這兩種生物互動關係常在野外同時存在，而它們的交互作用仍不清楚；第二是過去的研究通常沒有控制物種以及數量，所以難以釐清暖化影響生物群聚的機制。為了回答這兩個議題，本研究藉由操控性實驗探討暖化會如何影響1)螞蟻與蚜蟲的互利共生關係，2)瓢蟲與蚜蟲的掠食關係，和3)上述兩種關係的交互作用。本研究利用在台灣野外具互動關係的大豆、大豆蚜、七星瓢蟲及熱帶火蟻，進行2 x 2 x 3複因子設計實驗: 包含2組螞蟻處理(有/無) x　2組瓢蟲處理(有/無) x 3組溫度處理(控制組/+3ºC/+6ºC)，以上每組皆含大豆及蚜蟲。本研究的物種密度與控制組溫度均依據野外調查資料來設定，而暖化的增溫程度則依據聯合國氣候變遷委員會的預測來設定。本研究記錄蚜蟲數量、植株上的螞蟻數量以及瓢蟲的行為反應。結果顯示：一、 在控制組溫度下，螞蟻與蚜蟲的互利共生關係增加了264% − 328% 的蚜蟲數量，但這樣的效益卻隨著暖化而減弱。二、在控制組溫度下，瓢蟲與蚜蟲的掠食關係減少了38% 的蚜蟲數量。在暖化情況下，當有螞蟻存在時，瓢蟲的掠食關係會減少60% 的蚜蟲數量；然而，當螞蟻不存在時，瓢蟲掠食關係對蚜蟲數量的影響則會消失，雖然暖化下的瓢蟲掠食行為有提高的現象。三、 瓢蟲的存在會提升在植株上巡邏的螞蟻數量（+39%），暗示螞蟻為保護蚜蟲而與瓢蟲產生敵對的關係；此外，暖化似乎增強了這樣的敵對關係、改變瓢蟲偏好的微棲地、增加瓢蟲在敵對情況下的死亡率。本研究顯示暖化可減弱互利共生、減弱掠食關係及影響其交互作用，鑒於這些生物互動關係會影響生態系統的穩定，暖化有可能會透過改變這些關係進而降低生態系統的穩定性，相關的議題值得後續研究來驗證。

Mutualism (e.g., ants and aphids) and predation (e.g., ladybugs and aphids) are important species interactions that can shape community structure and stabilize ecosystems. As climate change proceeds, studies have investigated warming impact on these species interactions. However, two knowledge gaps exist. First, mutualism and predation were usually examined in isolation, although they often coexist. Therefore, their interaction remains to be answered. Second, previous studies usually did not explicitly manipulate species identities and densities. Therefore, the underlying mechanisms for warming impact may be confounded. To fill the knowledge gaps, this laboratory empirical study examined warming impact on 1) the mutualism between ants and aphids, 2) the predation of aphids by ladybugs, and 3) the interaction between mutualism and predation (e.g., antagonism). This study focused on the ant-aphid-ladybug system in Taiwan’s soybean farms, including the tropical fire ants (Solenopsis geminata), soybean aphids (Aphis glycines), and seven-spotted ladybugs (Coccinella septempunctata) on soybean plants (Glycine max). The experiment had a 2 x 2 x 3 factorial design: mutualism treatment (ant presence/absence) x predator treatment (ladybug presence/absence) x temperature treatment (control, +3, +6°C). Each treatment combination was performed with a caged soybean plant and aphid colony. The species densities and control temperature were based on field data, and the warming scenarios were based on the prediction by Intergovernmental Panel on Climate Change (IPCC). We recorded aphid density, ant density, and ladybug behavior. The results included the followings: 1) Ant-aphid mutualism increased aphid populations by 264% − 328% at control temperature, but this benefit reduced under warming. 2) Predation of aphids by ladybugs reduced aphid population by 38% at control temperature; this top-down control on aphid populations under warming reduced aphid population by 60% at ant presence but disappeared at ant absence, although warming increased the predation behavior of ladybugs. 3) Ladybug presence increased the number of patrolling ants (protecting aphids) on soybean plants by 39% on average, suggesting an antagonism between ants and ladybugs (i.e., interaction between mutualism and predation).Furthermore, warming seemed to intensity this antagonism, affect ladybugs’ location, and increase ladybugs’ mortality. Taken together, this study demonstrates that warming may reduce mutualism effect, reduce predation effect, and affect their interaction. Given that mutualism and predation can stabilize ecosystems, climate warming impact on these critical species interactions may potentially destabilize ecosystems and deserves further investigations.

口試委員審定書 i
中文摘要 ii
英文摘要 Abstract i ii
Introduction 1
Material and methods 7
Species 7
Experimental design 8
Experimental device 9
Ladybug’s behavior 9
Statistical analyses 10
Results 12
Warming impact on the mutualism between ants and aphids 12
Warming impact on the predation of aphids by ladybugs 13
Warming impact on mutualism-predation interactions 14
Discussion 16
Why warming reduced mutualism effect 17
Why warming reduced predation effect at ant absence, but increased at ant presence 19
How warming shifted the antagonism between ants and ladybugs 20
What value this study design provides 22
Which caveat this study comes with 22
Conclusions 23
Reference 25
附錄 30

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