本研究目的為瞭解關子嶺地區螢火蟲多樣性與大端黑螢 (Luciola anceyi) 之交配行為。野外實驗時間由2010年9月至2012年4月止。夜間螢火蟲調查為每週至關子嶺九龍山一號及石雅谷穿越線調查日落後一小時內螢火蟲物種及數量。日間螢火蟲調查由2011年3月至5月為每二週至此二條穿越線調查螢火蟲物種。另於冬季及秋季在二條穿越線取樣記錄植物種類及分層。另外，在九龍山二號、九龍山三號以及大凍山穿越線進行大端黑螢交配行為觀察，利用標記釋放探討大端黑螢雌雄蟲在日夜間之再捕捉率，並以數位錄影記錄分析大端黑螢交配過程及各行為間隔時間。
日夜間螢火蟲調查從2010年9月至2011年12月在關子嶺共發現6屬14種螢火蟲。日夜間螢火蟲於5月至6月最高各可記錄到8種，而1月、2月及9月則未有螢火蟲記錄。日間活動螢火蟲有4種，包括奧氏弩螢 (Drilaster olivieri) 、紅胸窗螢 (Pyrocoelia formosana) 、黑腹櫛角螢 (Vesta scutellonigra) 以及大端黑螢。夜間螢火蟲之Shannon-Wiener多樣性指數及均勻度於二條穿越線皆以夏季最高，顯著高於其他三個季節 (Tukey’s HSD, P＜0.05) 。春夏季日夜間螢火蟲記錄最高為12種，而秋冬季卻僅記錄4種螢火蟲。
2011年夜間螢火蟲物種相對頻度以黑翅螢 (Luciola cerata) 最高占68.21％，其次為端黑螢 (Luciola praeusta) 占9.48％、大端黑螢占6.42％、黃肩脈翅螢 (Curtos mundulus) 占5.92％、橙螢 (Diaphanes citrinus) 占4.47％以及山窗螢 (Pyrocoelia praetexta) 占3.29％，而其餘6種螢火蟲比例皆不到3％。黑翅螢於4月至5月日落後一小時平均密度為最高，在九龍山一號分別為22.77隻/ 100 m2及15.54隻/ 100 m2，而在石雅谷則為6.07隻/ 100 m2及5.91隻/ 100 m2。黑翅螢的出現數量與石雅谷溫度具有正相關 (P＜0.05) ，而大端黑螢的數量則與石雅谷的相對濕度具有負相關 (P＜0.001) 。端黑螢的數量在二條穿越線皆與溫度呈顯著正相關 (P＜0.05) ，而黃肩脈翅螢出現數量皆與石雅谷的溫度及相對濕度具有正相關 (P＜0.05) 。但橙螢與山窗螢的數量在二條穿越線均與溫度及相對濕度並無相關性 (P＞0.05) 。
植物分層在石雅谷較九龍山一號為複雜，會影響到螢火蟲群聚組成。石雅谷穿越線植物分層以喬木及草本二層 (冬季：31.82％；春季：33.64％) 與喬木、灌木及草本三層為最高 (冬季：30.91％；春季：34.55％) 。九龍山一號穿越線植物分層以單層草本比例最高 (冬季：47.50％；春季：48.75％) ，但卻缺乏單層灌木、喬木及灌木二層以及喬木、灌木及草本三層。黑翅螢偏好出現在九龍山一號的草原地，而橙螢及山窗螢則偏好出現在植物分層較複雜的石雅谷。
大端黑螢在日夜間皆具有求偶及交尾行為。在日間捕捉大端黑螢之飛行個體中，雄蟲數量顯著高於雌蟲 (P＜0.05) 。在夜間可再捕捉日間標記之大端黑螢雌雄蟲個體，但雄蟲再捕捉率為雌蟲的2倍 (分別15.94％與8.55％) ，此顯示成蟲在日間活動，同時在夜間於原穿越線活動發光。大端黑螢成蟲在日夜間皆表現交配行為，而雄蟲在日間可能以化學訊號作為尋找配偶。大端黑螢的交配行為無法對應目前已知螢火蟲的兩性溝通模式，因此可視為全新的活動類型及兩性溝通模式。
關子嶺地區雖有長年耕作與開發過度的影響，但螢火蟲數量及物種仍相當豐富且一年四季皆有螢火蟲的出現，因此關子嶺具有發展生態旅遊及賞螢的潛能。但賞螢也會增加環境的破壞，因此如何適度發展生態旅遊又不破壞棲地，考驗著當地永續經營的目標。期盼本研究結果能提供關子嶺之螢火蟲棲地保育及經營管理參考依據。

The objective of this study was to investigate the diversity of fireflies and the mating behavior of Luciola anceyi in Guanziling, Tainan. The field study was carried out from September 2010 to April 2012. The nocturnal survey of number of individuals and species of firefly was within one hour after sunset weekly in two transects, Jiulongshan 1 (JS1) and Shiya valley (SV). The diurnal firefly survey was from 2011 March to May once for every two weeks in these two transects. In addition, vegetation species and layers was recorded in these two transects during winter and spring. The sex ratio, population size and mating behavior of L. anceyi were studied in three transects (Jiulongshan 2, Jiulongshan 3 and Dadongshan) with mark-recapture and digital video camera.
Total 6 genus and 14 species of firefly were found in JS1 and SV, Guanziling from September 2010 to December 2011. The maximum number of firefly species reached its peak of 8 in May and June in Guanziling, but no adult firefly was found in January, February and September. Four species of fireflies were found active during daytime, including Drilaster olivieri, Pyrocoelia formosana, Vesta scutellonigra and L. anceyi . The Shannon-Wiener’s diversity index and evenness of nocturnal fireflies were both highest in summer, significantly higher than those from other seasons (Tukey's HSD, P＜0.05). During spring and summer, 12 species of firefly species were recorded but only 4 species was recorded during autumn and winter.
In 2011, Luciola cerata was the most dominant species in firefly community in Guanziling, accounted for 68.21%. The second comeon one was Luciola praeusta (9.48 %), followed by Luciola anceyi (6.42%), Curtos mundulus (5.92%), Diaphanes citrinus (4.47%), and Pyrocoelia praetexta (3.29%). The highest average density within one hour after sunset of L. cerata was from April to May (22.77/ 100 m2 and 15.54/ 100 m2 in JS1 and 6.07/ 100 m2 and 5.91/ 100 m2 in SV). The number of L. cerata was positively correlated with temperature in SV (P＜0.05), but the number of L. anceyi was negatively correlated with relative humidity in SV (P＜0.001). The number of L. praeusta was positively correlated with temperature of these two transects (P＜0.05), and C. mundulus was positively correlated with temperature and relative humidity (P＜0.05) in SV. But the number of D. citrinus and P. praetexta were neither correlated with temperature nor relative humidity from these two transects (P＞0.05).
Vegetation stratification in SV was more complex than in JS1, and firefly communities were influenced by plant stratification. The relative frequency of plant layer was the highest in two layers of shrub and herb (winter：31.82%；spring：33.64%) and three layers of tree, shrub and herb (winter：30.91%；spring：34.55%) among seven layers of plants in SV. On the other hand, The relative frequency was the highest in the single herb layer in JS1 (winter：47.50%；spring：48.75 %), and lacking herb layer, two layers of tree and shrub and three layers of tree, shrub and herb. L. cerata prefers to appear in the grasslands of JS1, and D. citrinus and P. praetexta prefer to appear in the habitat with complex vegetation in SV.
The courtship and copulation behavior of L. anceyi were recorded during daytime as well as nighttime. The number of adult male L. anceyi captured during daytime flying was significantly higher than adult female captured (P＜0.05). During nighttime, the recapture rate of marked adult males of L. anceyi was twice that of adult females (15.94 % versus 8.55 %). It indicates that the adult L. anceyi appears to be active in the daytime and nighttime in the same area. The mating behavior of L. anceyi occurrs both day and night, therefore, male L. anceyi most likely uses chemical signals to find mates during day time. The mating pattern of L. anceyi is not corresponded with any sexual communication systems known for fireflies. Therefore it can be regarded as a new type of activity and sexual communication systems for fireflies.
Although Guanziling area is influenced by long-term cultivation and over development, the quantity and species number of fireflies are plentiful and fireflies occur in all four seasons. Therefore Guanziling has the potential for developing ecotourism and firefly-watch. However, firefly-watch can also increase the damage to the environment. How to appropriately develop ecotourism without destroying the habitat may become the challenge and test for the goal of sustainable management. Finally, the results of this study can provide useful information for habitat conservation and management of fireflies in Guanziling.

論文審定書………………………………………………………………………………i
誌謝……………………………………………………………………………………...ii
中文摘要……………………………………………………………….……………….iii
英文摘要………………………..……………………………………………………….v
目錄……...…………………………………………………………………………….viii
圖次……………………………………………………………………………………...x
表次…………………………………………………………………………………….xii
前言…………………………………………………………………………………...…1
材料與方法……………………………………………………………………………...4
一、螢火蟲調查……………………………………………………………………….4
(一) 研究地點……………………………………………………………….………4
(二) 溫度、相對濕度及光照度測量………………………………………………...4
(三) 植被調查……………………………………………………………………….5
(四) 螢火蟲夜間物種調查……………….................................................................5
(五) 螢火蟲日間物種調查………………………………………………………….6
二、大端黑螢之族群及行為觀察…………………………………………………….6
(一) 研究地點……………………………………………………………………….6
(二) 大端黑螢之標記釋放再捕捉………………………………………………….7
(三) 大端黑螢之行為觀察………………………………………………………….8
三、資料分析與統計…………………………………………………………….…10
(一) 螢火蟲在夜間物種數量分析………………………………………………...10
(二) 螢火蟲多樣性分析……………………………………………………...……10
(三) 統計分析……………………………………………………………………...11

結果…………………………………………………………………………………….12
一、溫度、相對濕度及植被之變異………………………………………………...12
二、螢火蟲多樣性…………………………………………………………………...14
三、六種螢火蟲在夜間族群數量與環境因子的關係…………………………..….17
四、大端黑螢族群數量及交配行為……………………………………………...…20
討論…………………………………………………………………………………….22
一、螢火蟲多樣性…………………………………………………………………...22
二、螢火蟲數量與環境因子的關係………………………………………………...23
三、大端黑螢族群數量及交配行為……………………………………………...…25
四、總結及後續研究………………………………………………………………...28
參考文獻………………………………………………………………………….……30
圖…………………………………………………………………………………….…34
表……………………………………………………………………………………….51
附錄一：研究地點照片………………………………………………………………...60
附錄二：大端黑螢形態觀察………………………………………………..…………61
附錄三：統計分析……………………………………………………………………...66
附錄四：本研究之大綱………………………………………………………………...71

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