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Larval Morphology of Twenty One Species and Bionomics of Fireflies (Coleoptera: Lampyridae) in Taiwan
digital vedio record system
|摘要:||本文描述台灣產螢科21種末齡幼蟲的形態，分別為雙色垂鬚螢(Stenocladius bicoloripes Pic)、大端黑螢 (Luciola anceyi Olivier)、黑翅螢(Luciola cerata Olivier)、黃緣螢(Luciola ficta Olivier)、端黑螢(Luciola gorhami Ritsema)、條背螢(Luciola substriata Gorham)、黃胸黑翅螢(Luciola sp.1)、邊褐端黑螢(Luciola sp.2)、梭德氏脈翅螢(Curtos sauteri (Olivier))、橙螢(Diaphanes citrinus Olivier)、蓬萊短角窗螢(Diaphanes formosus Olivier)、鋸角雪螢(Diaphanes lampyroides (Olivier))、台灣窗螢(Pyrocoelia analis (Fabricius))、紅胸窗螢(Pyrocoelia formosana Olivier)、山窗螢(Pyrocoelia praetexta Olivier)、赤腹窗螢(Pyrocoelia sanguiniventer (Olivier))、長胸窗螢(Pyrocoelia sp.1)、北方鋸角螢(Lucidina biplagiata (Motschulsky))、鹿野氏黑脈螢(Pristolycus kanoi Nakane)、雲南扁螢(Lamprigera yunnana (Fairmaire))及赤腹櫛角螢(Vesta impressicollis Fairmaire)，文中討論背中線、體色型態、尾足數量與排列及棲地等特性，並將結果作成種級檢索表與屬級圖索表。
選擇台灣窗螢、黑翅螢、大端黑螢、端黑螢及邊褐端黑螢等5種常見陸生螢火蟲，利用2種方法飼育，皆能完成生活環。以”扁蝸牛活體飼養法”飼養台灣窗螢幼蟲，幼蟲期249.69 ±49.27天，成蟲羽化率48 %。以”扁蝸牛碎肉飼養法”飼養黑翅螢、大端黑螢、端黑螢與邊褐端黑螢，幼蟲期分別為182.17 ±13.80天、158.17 ±16.12天、220.92 ±21.29天與196.81 ±47.15天，其成蟲羽化率分別為40 %、43 %、37 %及45 %。各生長期中以幼蟲期80.3 ±3.3 %最高，成蟲期7.9 ±2.0 %，蛹期7.3 ±2.8 %，卵期6.0 ±1.3 %。這些雌性幼蟲期皆較雄性幼蟲期為長。台灣窗螢幼蟲期具4-7齡，小端黑螢與邊褐端黑螢幼蟲期具4-6齡，大端黑螢幼蟲期具4-5齡，黑翅螢幼蟲期具5齡，在此就5種陸生螢火蟲生活環提出2類基本型式。這些特性表現生活史可塑性。台灣窗螢幼蟲不會建築土繭，可直接進入前蛹期；其餘種類末齡幼蟲會建築橢圓形土繭，且依種類而略有不同。5種螢火蟲幼蟲皆為肉食性，以台灣窗螢幼蟲食性較專一，其餘種類末齡幼蟲食性較複雜。
描述台灣窗螢雌蟲生殖系統，其卵巢屬於無營養細胞型。雌蟲單隻飼養在25-28℃室溫下，其成蟲壽命12.76 ±5.32天，產卵前期1.24 ±1.62天，產卵期5.08 ±3.18天，產卵後期 1.78 ±0.45天。交尾後第一天之產卵量最高，然後依次遞減 。雌蟲產卵量332 ±114粒，最高538粒，最低120粒。依據台灣窗螢產卵結果，提出載卵壓力假說。已交尾雌蟲產卵之卵徑1.61 ±0.07 mm，大於無交尾雌蟲所產卵之卵徑1.32 ±0.05 mm，兩者差異顯著。雌蛹重量與產卵量呈正相關。
描述與測量25種台灣產螢火蟲的卵，其中22種為圓球形，3種為橄欖球形。其中雲南扁螢卵徑2.38 mm最大，黃緣螢卵徑0.50 mm最小。以卵徑、雌蟲類型與雄蟲體長等資料建構資料矩陣，經聚類分析結果，可分為三群，A群以螢亞科螢火蟲與部份熠螢亞科種類為主，卵皆呈圓球形；其群可分為二亞群，I亞群有窗螢屬、短角窗螢屬與雲南扁螢等9種，具雌雄二態型，雌蟲小翅型，經相關分析結果雄蟲和雌蟲體長與卵大小間呈正相關(p < 0.01)；II亞群有鹿野氏黑脈螢、赤腹櫛角螢、紅胸黑翅螢與黑翅螢。B群以熠螢亞科為主，包括熠螢屬與脈翅螢屬之螢火蟲，除紋螢卵呈橄欖球形外，其餘卵皆球形，比較其中7種熠螢屬螢火蟲卵徑與體長，不具相關性；但其中以黃緣螢與黃胸黑翅螢二種水生螢火蟲卵徑明顯較小。C群為大場雌光螢與雙色垂鬚螢，卵呈橄欖球形，雌蟲屬蠕蟲型。在卵徑與產卵量間之關係上，水生的黃緣螢、黃胸黑翅螢總產卵量明顯地高於陸生端黑螢，在體長、卵徑與產卵量間呈權衡現象。雲南扁螢與大場雌光螢的蠕蟲型雌蟲具有慈愛行為，所產的卵有較大的趨勢，本研究提出3類螢火蟲卵徑趨勢，並討論在演化與生態上的關係。
本研究自行規劃組裝之數位錄影系統，包括電腦主機系統、影像擷取卡、40G硬碟、12倍速光碟燒錄機、高畫質監視器CCD、紅外線燈源、影像作業軟體與相關周邊設備。本系統與錄影帶錄影系統和DV攝影系統相較下，其功能、資料管理、速度與可擴充性皆較佳，能夠有效地在夜間工作，適合作為夜行性螢火蟲活動的監視與記錄系統。在化蛹與羽化之發生頻度呈逢機分布，雌蟲化蛹平均所需時間58.37 ±26.89 分鐘，而雄蟲為64.50 ±47.96分鐘。雌蟲羽化平均所需時間101.66 ±52.60分鐘，而雄蟲為85.04 ±57.96分鐘，兩者在雌雄之間無顯著差異(t-test, p > 0.05)。|
Twenty one species of firefly larvae in Taiwan were described and included with Stenocladius bicoloripes Pic, Pristolycus kanoi Nakane, Luciola ficta Olivier, Luciola sp.1, Luciola substriata Gorham, Luciola anceyi Olivier, Luciola cerata Olivier, Luciola gorhami Ritsema, Luciola sp.2, Curtos sauteri (Olivier), Diaphanes citrinus Olivier, Diaphanes formosus Olivier, Diaphanes lampyroides (Olivier), Pyrocoelia analis (Fabricius), Pyrocoelia formosana (Olivier), Pyrocoelia praetexta (Olivier), Pyrocoelia sanguiniventer (Olivier), Pyrocoelia sp.1, Lucidina biplagiata (Motschulsky), Lamprigera yunnana (Fairmaire), Vesta impressicollis Fairmaire. The characteristics of the central line on the dorsal plate, body color pattern, number and arrangement of proleg, and habitats were discussed. The results presented the key to the common species in Taiwan. This paper presents rearing methods for the five species of the common terrestrial fireflies, Pyrocoelia analis, Luciola cerata, Luiola anceyi, Luciola gorhami, and Luciola sp. and completed their life cycles. The larval duration and emergence ratio show 249.69 ±49.27days (mean ±SD) and 48 % brood by “living snail rearing method” for P. analis. The larval duration of terrestrial larvae, L. cerata, L. anceyi, L. gorhami, and L. sp. brood by using “snail meat rearing method” show 182.17 ±13.80 days, 158.17 ±16.12 days, 220.92 ±21.29 days and 196.81 ±47.15 days and emergence ratio show 40 %, 43 %, 37 %, and 45% respectively. According to the results for the life cycles of five species fireflies, we proposed two basic types of life cycle. Ratio in the different life stage could find the highest 80.3 ±3.3 % of larval duration, 7.9 ±2.0 % of adult duration, 7.3 ±2.8 % of pupa duration and 6.0 ±1.3 % of egg duration. Their female larval duration is longer than males. The larval stage consists of 4th- 7th instars in P. analis, 4th- 6th instars in L. gorhami and L. sp., 4th- 5th instars in L. anceyi and 5th instars in L. cerata. These traits show the life-history plasticity. The last instar larvae of P. analis did not build the mud cocoon and ran into prepupa stage directly. The others would build the mud cocoon and shape varies among different species. All larvae are carnivorous. The larvae of P. analis has a more specific diet than the other species. This paper describes the reproductive system of the firefly, Pyrocoelia analis (Fabricius). Its ovarioles belong to the panoistic type. In the laboratory at 25-28℃, its longevity was 12.76 ±5.32 (mean ±SD) days, and preoviposition, oviposition, and postoviposition periods of the female were, 1.24 ±1.62, 5.08 ±3.18 and 1.78±0.45 days, respectively. Most of the eggs were deposited on the same day as mating, and daily egg production decreased successively. The total number of eggs produced per female was 332 ±114 with maximum of 538 and minimum of 120. We propose the egg-loaded pressure hypothesis as an interpretation of this oviposition behavior. Egg sizes of mated females were 1.61 ±0.07 mm, which were significantly larger than the 1.32 ±0.05 mm of unmated females. The female pupa weight was significantly positively related to egg production. Twenty-two species for ball shape and three species for ellipsoid shape of eggs was found 25 species of the fireflies in Taiwan The largest egg size of firefly was for the firefly Lamprigera yunnana (Fairmaire) which was 2.38 ±0.08 mm (mean ±SD) while the smallest was the firefly Luciola ficta which was 0.50 ±0.06 mm. According to the clustering analysis on characters of eggs size, female type and male length, their dendrogram was constructed. They were divided three groups. The A group included all the Lampyrinae species and a part of Luciolinae species, which had the ball shape eggs. The group was subdivided two subgroups. Subgroup I included nine species belonging to the genera Pyrocoelia, Diaphanes and Lamprigera. All females show sexual dimorphism and the brachyptery. Their egg sizes and body lengths of male and female were significantly positive correlated. Subgroup II included Pristolycus kanoi Nakane, Vesta impressicollis Fairmaire, Luciola cerata Olivier and Luciola kagiana Matsumura. The B group included Luciola spp. and Curtos spp. of Luciolinae. Their eggs were ball shape, except Lucola filiformis Olivier which had the ellipsoid shape egg. Their egg sizes and body length in seven species were not significantly correlated. The egg sizes of the aquatic fireflies, Luciola ficta Olivier and Luciola sp.1 were significantly smaller than the terrestrial species. The C group included Stenocladius bicoloripes Pic and Rhagophthalmus ohbai Wittmer which had ellipsoid shape eggs deposited by worm-liked females. The fecundity of aquatic fireflies, Luciola ficta Olivier and Luciola sp.1 is higher than the terrestrial firefly, Luciola gorhami Ritsema. In the respects of body length, egg size and fecundity could find trade-off. The egg sizes with maternal care deposited by the worm-liked females, L. yunnana and R. ohbai showed the larger size trend. Three types of trends in egg sizes of fireflies were proposed and discussed to the evolutional and ecological relations. We set up a digital video system that connected to the main system of personal computer with an image card, a 40 G hard disk, a 12 X CDR, and a high quality CCD. It was provided with infrared light as its illumination source. This system was found to be better than the video system and the digital video camera system in functional quality and efficiencies, such as image view, storage, transfer and extension, and data collection and management. The digital video system is a useful monitoring system particularly suitable for studies on activities of nocturnal fireflies. In this study the digital video system was tested to monitor the pupation and emergence of the firefly Pyrocoelia analis Fabricius. The results showed that occurrence of the pupation and emergency during 24 hours of the day were the random phenomena without a peak occurrence. The time duration for the pupation was 58.37 ±26.89 min for females and 64.50 ±47.96 min for males, while for the emergence was 101.66 ±52.60 min for females, and 85.04 ±57.96 min for males. However, there was no sexual significant difference in these two characters (t-test, p > 0.05).
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