本研究利用的物種為綠島龜鼻灣海草床中的一種小型螺類項鍊蟹守螺 (Cerithium zonatum)，由於其數量多且能利用齒舌刮食植物產生特殊的聲音，因此可以利用其聲音訊息來研究其攝食活動，作為海草生態系中優勢底棲無脊椎動物的行為生態的參考資料。本實驗於2007 年11~12 月、2008 年3~6 月和2009年4 月~7 月間共十個月在綠島龜鼻灣及紫坪兩地進行。利用其對三種食性的偏好程度不同探討其在三種微棲地 (在海草、海藻及附著微細藻的石頭上)的分佈。並根據其日夜週期及其高低水溫的變化探討不同環境因子對其攝食活動影響。亦探討大小個體在攝食活動之間是否有所不同。也利用電子顯微鏡研究其齒舌的形式。結果顯示，項鍊蟹守螺在海草上的刮食頻度比較高，此可能為蟹守螺選擇海草為主要棲地，而此推論與在三個微棲地採樣的項鍊蟹守螺數量密度的結果符合。而強日光與高水溫確實會提升刮食頻度，而黑暗無光線與低水溫會使其攝食活動停滯。小個體的刮食頻度比大個體多，但是在齒舌上面沒有差異，推測可能是小個體需要多攝食以用於個體的成長。最後也探討漁舟蜑螺 (Neritaalbicilla)的刮食聲音特性，其脈衝波數多於項鍊蟹守螺，推測可能是齒舌構造不同所致，而且項鍊蟹守螺刮食效率比漁舟蜑螺佳。由本研究結果可以得知利用聲音訊息探討小型螺類攝食活動是可行的，並期望能應用到其他使用齒舌刮食的軟體動物上。

This study focused on a small gastropoda, Cerithium zonatum, found in seagrass meadow at Gui Wan Bi, Green Island, Taiwan. Because this species is high in density in the seagrass meadow and it uses its specific radular to scrap the substrate and emits
a particular sound, we can use this acoustic clue to study its feeding activity. C.zonatum samples were collected from Gu Wan Bi, and Zi-Ping Green Island between November to December 2007,March to June 2008, and April to July 2009. It distributes in three habitats (i.e. seagrass、macro algae、rubble) and scrapping sounds were recorded while they were in these habitats. Light and water temperature were
treated as experimented factors and their influence on the feeding frequency was deciphered. I also compared individuals of different sizes to find out the differences in feeding frequency between them. SEM was used to observe the radular of C. zonatum. According to the results, I infer that C. zonatum might prefer seagrass meadow as its must suitable microhabitat where feeding activity was higher. Sunlight and high water temperature might lead to increases in feeding activity, but darkness
and low water temperature showed opposite effect; it stopped feeding when water temperature had dropped to 17℃. Scrap frequency of small individuals was higher than larger ones, and their radular were similar in their morphology. I infer that small
ones invest more time and energy in feeding in order to grow. Scrapping sound of C.zonatum was compared to Nerita albicilla; there were pulses in the sound of N.albicilla than C. zonatum. It is inferred that this difference might be due to the difference between their radular morphology. I consider that using an bioacoustic approach to study the feeding activity of small gastropoda is feasible.

目錄
章次 頁數
謝辭....................................................................................... Ι
中文摘要.............................................................................. Ⅱ
英文摘要.................................................................. ...........Ⅲ
壹、前言..................................................................... .........1
1.海草生態與黑潮.............................................................. 1
2 植食性螺類的攝食機制................................................... 2
3.植食性腹足綱動物齒舌形式......................................... ..2
4.利用動物聲學在無脊椎動物的研究.............................. 3
5.研究物種..................................................... .....................4
6.研究目的............................................................... ...........5
貳、材料與方法......................................................... ........6
1.採集地點................................................................ .........6
2.錄音系統......................................................................... 6
3.項鍊蟹守螺刮食聲音訊息參數.............................. .......6
4.以食物選擇推論三個微棲地不同數量密度的分佈.. ...6
4-1.數量密度的調查.................................. .......................6
4-2.攝食的投資-三種食物的測試..................................... 7
5.攝食活動的晝夜變異..................................................... 7
5-1.二十四小時的攝食活動變化..................................... 7
5-2.黑暗在白天對刮食活動的影響................................. 7
5-3.光線在夜間對刮食活動的影響................................. 7
6.溫度影響攝食活動頻率................................................ 8
6-1.觀察在不同水溫的活動情況..................................... 8
6-2.高低水溫對其攝食活動的影響................................. 8
7.大小不同的蟹守螺攝食活動頻率................................. 8
8.齒舌的研究..................................................................... 9
9.統計方法......................................................................... 9
參、結果.......................................................................... 10
1.項鍊蟹守螺的刮食聲音特徵...................... 10
2.以食物選擇推論三個微棲地不同數量密度的分佈.... 10
2-1.數量密度的調查........................................................ 10
2-2.攝食的投資-三種食物的測試................................... 10
3.日夜週期影響攝食活動................................................ 11
3-1.二十四小時的攝食活動變化..................................... 11
3-2.黑暗在白天對刮食活動的影響................................. 12
3-3.光線在夜間對刮食活動的影..................................... 13
4.溫度變化影響攝食活動................................................. 14
5.個體大小攝食活動......................................................... 16
6.齒舌形態......................................................................... 17
肆、討論............................................................................ 18
1.由食物選擇推論三個微棲地中項鍊蟹守螺密度的分
佈.18
2.日夜週期影響攝食活動................................................ 19
3.溫度變化影響攝食活動................................................ 20
4.個體大小攝食活動與比較齒舌差................................ 21
5. 齒舌型式與刮食聲音訊息的關係............................... 21
伍、結論........................................................................... 22
陸、參考文獻................................................................... 23
個人履歷…………......................………………………. 48
表目錄 頁數
表一、 實驗項目明細...................................................... 26
表二、使用One-way ANOVA 及Fisher's LSD 比較項鍊
蟹守螺在三種食物...............................................27
表三、使用隨機完全區集設計及Fisher'sLSD 比較項鍊
蟹守螺在正常日間及給予遮光下的攝食活動..28
表四、使用隨機完全區集設計及Fisher'sLSD 比較項鍊
蟹守螺在正常夜間及照光下的攝食活動...........29
表五、使用隨機完全區集設計及Fisher's LSD 比較項鍊
蟹守螺在高水溫下的攝食活動...........................30
表六、使用隨機完全區集設計及Fisher's LSD 比較項鍊
蟹守螺在低水溫下的攝食活動............................31
表七、使用隨機完全區集設計及Fisher's LSD 比較不同
殼長項鍊蟹守螺的攝食活動................................32
圖目錄 頁數
圖一、 實驗研究地點:台灣台東縣綠島鄉.................... 33
圖二、 項鍊蟹守螺 (Cerithium zonatum)標本...............34
圖三、 項鍊蟹守螺刮食聲音參數.................................... 35
圖四、 項鍊蟹守螺 (Cerithium zonatum)在綠島龜鼻灣海
草床的三個微棲地分佈圖.....................................36
圖五、 項鍊蟹守螺在泰來草上的刮食聲音分析圖........ 37
圖六、 項鍊蟹守螺在泰來草上的單一個刮食聲音分析
圖.............................................................................38
圖七、 2007~2009 年於三個微棲地中項鍊蟹守螺的密度
比較........................................................................ 39
圖八、 綠島龜鼻灣與紫坪兩地的採集項鍊蟹守螺殼之殼
長分佈................................................................. .....40
圖九、 項鍊蟹守螺刮食每分鐘刮食次數之晝夜分佈.....41
圖十、 項鍊蟹守螺的齒舌在解剖顯微鏡下的影像........42
圖十一、 項鍊蟹守螺的齒舌在電子顯微鏡下的影像... 43
圖十二、 項鍊蟹守螺的單一排齒舌的放大對照圖....... 44
圖十三、 項鍊蟹守螺大小殼長的齒舌比較圖................45
圖十四、 漁舟蜑螺 (Nerita albicilla)標本......................46
圖十五、 漁舟蜑螺 (Nerita albicilla)刮食聲音分析圖... 47

任淑仙，2000，無脊椎動物學。淑馨出版社，113 頁。
邵廣昭，1998，海洋生態學。文海環境科學叢書，210 頁。
柯智仁，2004，台灣海草分類與分布之研究。國立中山大學生物科學系碩士論文，
第10 頁。
張景平，黄小平，2008，海草與其附生藻類之間的相互作用。生態學雜誌27：
1785-1790。
賴景陽，2005，台灣貝類圖鑑。貓頭鷹出版社，155 頁。
賴亦德，陳俊宏，2004，探討光潤金線蛭 (Whitmania laevis) 捕食有口蓋淡水
螺類之偏好。特有生物研究，67-78頁。
Aguzzi, J. 2004. Feeding activity rhythm of Nephrops norvegicus of the western
Mediterranean shelf and slope grounds. Marine Biology 144: 463–472.
Akamatsu, T., Okumura, T., Novarini, N., Yan, H. 2002. Empiricalre finements
applicable to the recording of fish sounds in small tanks. The Journal of the
Acoustical Society of America. 112:3073–3082.
Au, W. W. L., Banks, K. 1998. The acoustics of the snapping shrimp Synalpheus
parneomeris in Kaneohe Bay. The Journal of the Acoustical Society of
America 103:41–47.
Barnes, B.M. 2000. Effects of ambient temperature on metabolic rate, respiratory quotient,
and torpor in an arctic hibernator. American Journal of Physiology 279: 255-262.
Charnov, E. L. 1976. Optimal foraging, the marginal value theorem. Theoretical Biology
9:129-136.
Duarte, C.M. 1999. Seagrass biomass and production: a reassessment. Aquatic Botany,
65:159–174.
Duffy, J.E. 1998. The Central Role of Grazing in Seagrass Ecology, Seagrass: Biology,
Ecology and Conservation 463-501.
Houbrick, R. S. 1992. Monograph of the Genus Cerithium Bruguiere in the Indo-Pacific
(Cerithiidae: Prosobranchia). Smithsonian Contributions to Zoology 510:211-223.
Hays, C. G. 2005. Effect of nutrient availability, grazer assemblage and seagrass source
population on the interaction between Thalassia testudinum (turtle grass) and its algal
epiphytes. Journal of Experimental Marine Biology and Ecology 314:53– 68.
Hauxwell, J. 2003. Eelgrass Zostera marina loss in temperate estuaries: relationship to
land-derived nitrogen loads and effect of light limitation imposed by algae. Marine
Ecology Progress Series 247: 59–73.
Heck, K. L. Jr. 2006. Plant–herbivore interactions in seagrass meadows. Journal of
Experimental Marine Biology and Ecology 330:420-436.
Harley, E. H., J. W. Alan. 1996. Dynamics of the Kuroshio/Oyashio current system using
eddy-resolving models of the North Pacific Ocean. Journal of geophysical research
101:941-976.
Kitting, C.L. 1979. The Use of feeding noises to determine the alga foods being
consumed by individual Intertidal Molluscs. Oecologia 40:1-17.
Levinton, J. S. 1995. Marine biology: Function, biodiversity, ecology. Oxford University
Press, New York 22:451-462.
McGlathery, K. J. 1995. Nutrient and grazing influences on a subtropical seagrass
community. Marine Ecology 122: 239-252.
Michael, A. B. 1998. Epiphytes of Seagrasses, Seagrass: Biology, Ecology and
Conservation 441-461.
Norton, T. A. 1990. Scraping a living: a review of littorinid grazing. Hydrobiologia 193:
117-138.
Orth, R. J., Carruthers, T. J. B., Dennison, W. C., Duarte, C. M., Fourqurean, J. W. 2006.
A Global Crisis for Seagrass Ecosystems. BioScience 56:987-996.
Pittendrigh, C . S. 1981. Circadian Systems: Entrainment. In Handbook of behavioral
neurobiology Biological Rhythms 4:57-80.
Patek, S. N. 2001. Spiny lobsters stick and slip to make sound. Nature 411:153-154.
Pechenik, J. A. 2000. Biology of the invertebrates. Fourth edition. McGraw-Hill,
Singapore.
Runham, N. W., Thornton, P. R., Shaw, D. A., Wayte, R. C. 1969. The mineralization and
hardness of the radular teeth of the limpet Patella vulgata L. Cell and Tissue Research
99:608-626.
Rossetti, Y., Cabanac, M. 2006. Light versus temperature: An intersensitivity conflict in
a gastropod (Lymnaea auricularia). Journal of Thermal Biology 31:514-520.
Sih, A., Christensen, B. 2001. Optimal diet theory: when does it work, and when
and why does it fail? Animal behavior 61:379-390.
Sand, J. K. 1989. Environmental variables and their effect on photosynthesis of
aquatic plant communities. Aquatic Botany. 34:5-25.
Symes, M. 2007. Subsurface Noise. Nature 19: 50-53.
Sommer, J. R. 1976. The ultrastructure of the mammalian cardiac muscle cell-with special
emphasis on the tubular membrane systems. A review. The American Journal of
Pathology 82: 192–232.
Scheltema, A. H., Kerth, K. 2003. Original molluscan radula: Comparisons among
Aplacophora, Polyplacophora, Gastropoda, and the Cambrian fossil. Journal of
Morphology. 257:219-245.
Williams, S. L., Ruckelshaus, M. H. 1993. Effects of nitrogen availability and herbivory on
eelgrass (Zostera marina) and epiphytes. Ecology 74:904-918.
Wu, W.L. 1986. SEM Studies on Radulae of the Taiwan Molluscs. Bull. Inst. Zool.,
Academia Sinica Supplement 14: 91-100.