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研究生: 戴翊翎
Tai, Yik-Ling
論文名稱: 東亞摺翅蝠形態及生理變異與其外寄生蟲感染的關係
Ecomorphology, body conditions, and differential ectoparasitic infections in Eastern Bent-winged Bats
指導教授: 李亞夫
Lee, Ya-Fu
學位類別: 碩士
Master
系所名稱: 生物科學與科技學院 - 生命科學系
Department of Life Sciences
論文出版年: 2020
畢業學年度: 108
語文別: 英文
論文頁數: 87
中文關鍵詞: 蝙蝠蝠蠅體況東亞摺翅蝠蛛蠅科寄生蟲寄生蝠蠅科蝠蟎屬
外文關鍵詞: bats, bat flies, body conditions, Miniopterus fuliginosus, Nycteribiidae, parasites, parasitism, Spinturnix, Streblidae
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  • 蛛蠅與蝠蟎是蝙蝠身上典型常見的寄生蟲,具有相當高程度的宿主專一性。蝙蝠的身體狀況、生殖周期產生的生理條件改變、及季節性的氣候變化,對寄生蟲的感染模式均可能有不同程度的影響。本研究針對東亞摺翅蝠(Miniopterus fuliginosus Hodgson, 1835),探討包括宿主個體形態與身體狀況、生殖行為所產生的生理變化、以及季節性氣候變化等因子對其感染程度的影響力。我檢測假說宿主的體況、性別、與生殖狀態對寄生蟲感染有影響,且預測體況較好的蝙蝠能為寄生蟲提供較佳資源,或是有更高的耐受性,其感染率會因此較高。我亦預測雌蝠的寄生蟲感染率會因生殖的生理需求而比雄蝠較高,且生殖季期間的雌性蝙蝠寄生蟲感染率將會更高。此外,我檢測氣候因子變化,包括溫度、濕度、雨量等對寄生蟲豐富度的影響。
    本研究於2016年9月至2018年9月期間進行取樣,於恆春龜仔角實驗林地中捕抓東亞摺翅蝠,記錄牠們的年齡、性別、體重等體況、測量翅膀型值,並採集其外寄生蟲,共記錄包括蛛蠅科 (Nycteribiidae) 的姜宜蛛蠅 (Penicillidia jenynsii, Westwood, 1834)、長鋏蛛蠅 (Nycteribia allotopa, Sperser, 1901)、短鋏蛛蠅 (N. parvula, Sperser, 1901)、蝠蠅科 (Streblidae) 的Ascodipteron speiserianum (Muir, 1912) 等4種雙翅目寄生蟲、及蝠蟎科 (Spinturnicidae) 蝠蟎屬Spinturnix spp.。
    整體而言,寄生蟲呈現明顯的季節性變化,在寒冷的月份數量較少,在溫和的月份數量較多,與月平均溫度有正相關。蛛蠅雌性個體均比雄性多,且全年生育,其中姜宜蛛蠅的生殖率在蝙蝠生殖季期間也有所增加。短鋏蛛蠅與蝠蟎在宿主上的豐度最高,而A. speiserianum在宿主上的豐度最低,但生物量上則比率佔最高。東亞摺翅蝠的雄性體形比雌性略大,生殖周期在暖和的五至八月發生,生殖中的母蝠,尤其是哺乳中的母蝠不單體況值比其他蝙蝠高,身上的寄生蟲亦為最多。懷孕與否與感染率無顯著相關。蝙蝠的體況、性別、與生殖狀態對寄生蟲感染皆有影響,其中長鋏蛛蠅、短鋏蛛蠅、與蝠蟎呈現相類似的感染模式,均偏向感染體況較好的宿主與生殖周期的母蝠,姜宜蛛蠅明顯地全年顯示雌性宿主偏向,且不受生殖狀態影響,而A. speiserianum則只偏向體況較好的宿主。非生殖季期間,寄生蟲感染率在公、母蝠之間並沒有顯著差異,亦不受公蝠的生殖狀態 (睪丸腫大) 影響。
    本研究結果顯示寄生蟲豐富度的季節性變化主要因宿主生殖期的生理變化以及相應的體況改變,受氣候因子的影響則相對較低。寄生蟲種之間的相關性因種間的形態、生活史、習性、生物量等不同而有所差異。母蝠的高感染率主要成因為生殖周期間的生理改變,並非全年性的現象。懷孕母蝠與哺乳母蝠之間的感染率差異亦反映了寄生蟲除了受宿主免疫系統影響,其他因子如宿主間的接觸、理毛等去除寄生蟲的行為對宿主間的感染率差異影響也許更大。因體況較好的宿主可給予寄生蟲的潛在能量較高,體況較好的宿主對寄生蟲可能有更高的耐受性,感染體況好的宿主比感染抵抗力較差但體況差的宿主對寄生蟲也許更有優勢。整體而言,本研究區別宿主體況、形態、生殖狀況、氣候因子等因子對東亞摺翅蝠外寄生蟲感染的影響,以釐清外寄生蟲在東亞摺翅蝠的感染模式,有助進一步了解群居食蟲性蝙蝠的宿主-寄生蟲關係,及其對不同族群組成之宿主的可能效應。

    Bat flies (Nycteribiidae and Streblidae) are ectoparasites of bats and often form a strong association and certain specificity with their hosts. Both host characteristics, in conjunction with seasonal breeding cycle of bats, and environmental factors may contribute to shape this rather unique host-parasite relationship and influence their infection. This study investigated the infection of Eastern Bend-winged Bat (Miniopterus fuliginosus, Hodgson, 1835) by ectoparasites. I examined effects of ecomorphology and body conditions of hosts on the extent of infection, and the seasonal distribution pattern of ectoparasites within population. I tested the hypothesis that parasite loads of individual bats are influenced by host body conditions, sex, and reproductive states, and predicted that better conditioned hosts, as resource, will be more tolerant to parasites over poorer conditioned hosts. I also predicted that females will be more heavily infected than males because of their reproductive investment, particularly during breeding season. In addition, I also examined the effects of seasonal weather factors on parasite abundance.
    Bats were sampled monthly between 2016 and 2018 in Kenting, Taiwan. Each bat was examined for sex and reproductive status, distinguished between adults from juveniles, their body mass and flight-associated eco-morphological traits were measured, and ectoparasites were collected. This study focused on bat flies and identified three species of Nycteribiidae, Penicillidia jenynsii (Westwood, 1834), Nycteribia allotopa (Sperser, 1901), Nycteribia parvula (Sperser, 1901), and one species of Streblidae, Ascodipteron speiserianum (Muir, 1912). I additionally collected mites and identified Spinturnix spp. (Spinturnicidae) as constituting the dominant portion.
    Both bat flies and mites revealed a distinct seasonal pattern in parasite abundance, lower in the winter from November to February and higher between March and October. Among climate factors, only ambient temperature positively correlated with parasite load, but not humidity or precipitation. All nycteribiid flies had female-biased population ratios and reproduced all year, and P. jenynsii also increased reproduction during the breeding season of the bats. Nycteribia parvula and spinturnicid mites dominated other species in numbers. Ascodipteron speiserianum occurred in low abundance, but dominated in biomass. Bent-winged Bats exhibited male-biased size dimorphism, and body conditions positively correlated with parasite abundance. During the breeding period of bats, generally around May to August and coincided with the warmer months, reproductive females, especially lactating bats, had higher body conditions and were generally most infested. The differences in parasite infection between pregnant and non-reproductive females were not apparent. Nycteribia allotopa, N, parvula, and Spinturnix spp. shared similar infection patterns, showing bias towards both reproductive females and bats with high BCI values. Penicillidia jenynsii showed a consistent female-bias pattern, and infection of A. speiserianum was biased towards high host BCI values regardless of sex nor reproductive states of the bat hosts. Outside of breeding season, I found no differences in overall parasite load between male and female bats, and the reproductive status of males did not affect their parasite abundance.
    My results indicated that the seasonal pattern of ectoparasite infection on bats was mainly caused by the changes in host states during the breeding season, which also contributed to changes in body conditions of bats, whereas the effects of weather factors were limited in comparison. Parasite species of similar phenology, life histories, and host adaptations were associated with each other, whereas high parasite biomass was negatively correlated with species co-occurrence. Female-biased parasitism was mainly caused by the changes in reproductive states in bats, but did not apply in a general sense. My study also highlighted the differences in infection between pregnant and lactating bats, which suggested exposure to parasites and anti-parasitic behavior to be more influential in infection rates between reproductive female bats, than differences in immune systems. In addition, my results suggested that bat flies aggregated on host bats with good body conditions, indicating higher energy reserves, over hosts with poor body conditions, indicating weaker immune defense. This implies that hosts with energy and resources of higher quality can tolerate more parasites over those with vulnerable immune defense. By distinguishing the effects of climate factors, host body conditions, and ecomorphology on the variation of ectoparasite load on Eastern Bent-winged Bats, this study discussed the causes of seasonal parasite infection patterns, which may help provide more insights to the host-parasite relationships in bats.

    摘要 i ABSTRACT iii ACKNOWLEDGEMENTS vi TABLE OF CONTENTS vii LIST OF FIGURES ix LIST OF TABLES x INTRODUCTION 1 STATEMENT OF THE PROBLEM 1 BACKGROUND 3 Roles of age, sex, and reproductive status of hosts 4 Body conditions of hosts 6 Fluctuating asymmetry 8 Interactions within parasites assemblages 9 Bats and ectoparasites 10 Ectoparasites of bats in Taiwan 12 The Eastern Bent-winged Bat 13 OBJECTIVES 14 MATERIALS AND METHODS 17 Study area 17 Study animals 18 Bat sampling 19 Ecomorphological measurement 19 Ectoparasite sampling 20 Data analysis 20 RESULTS 24 Bat phenology 24 Body mass and ecomorphology of bats 24 Body conditions of bats 29 Composition of parasites 29 Temporal variations in parasite load and diversity 32 Sex and pregnancy ratios of bat flies 38 Associations between parasites 38 Effects of host body condition on parasite load 42 Effects of host reproductive status on parasite load 44 DISCUSSION 49 Temporal variations of parasite load and the effect of weather factors 49 Sex ratio and pregnancy in nycteribiid flies 52 Diversity and interspecific associations in parasite assemblages 53 Relationships of ecomorphology, individual states, and parasite load of bats 56 Conclusion 60 LITERATURES CITED 63 VITA 87

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