藤壺 (Crustacea：Cirripedia)幼生發育的研究多聚焦在淺海的物種。在深海中，僅有一些鵝頸藤壺 (Order Pedunculata)的幼生發育已經被描述，然而對笠藤壺 (Order Sessilia)幼生發育仍多未知。此外，淺海藤壺的幼生普遍存在趨光行為，但對於生活在光線有限的深海藤壺，牠們的幼生是否也具有趨光行為，仍然值得研究。薄殼條藤壺Striatobalanus tenuis (Hoek，1883)為分布在中國東海和南海的深海藤壺 (300 - 400m)，該物種附著在岩石、腹足類動物和蟹類之外殼表面，為台灣底拖漁業常見的混獲物。在本研究中，我們定期到蚵仔寮漁港收集薄殼條藤壺的成體，在20°C下培養牠們的受精卵塊，孵化後的幼生用微藻餵養，每週更換一次水，並頻繁去除可能干擾幼生活動的沉澱微藻。我們成功觀察到六個時期的無節幼生和一個時期的腺介幼生，且不同時期無節幼生的形態，符合藤壺“正常”發育特徵。本研究中，使用紅色、綠色、藍色和白色光來測試薄殼條藤壺無節幼生的趨光性。第二期和第三期的幼生被所有光照設置所吸引；第四期和第五期幼生則對藍光表現出負趨光性，對其他實驗光源仍表現出正趨光性；第六期幼生對所有實驗光源皆表現出負趨光性。不同無節幼生時期的光照偏好變化可能與牠們垂直遷移的順序有關。我們推測薄殼條藤壺的幼生在被母體釋放後會遷移到約100米深的淺海中，以攝食浮游植物，然後遷移回深海進行沉降。

Larval development of many shallow water barnacle species (Crustacea: Cirripedia) has been well studied. In the deep sea, the larval development of some goose barnacles (Order Pedunculata) was briefly described but remained unknown for acorn barnacles (Order Sessilia). In addition, the presence of phototactic behaviors, which are common in larvae of shallow water species, remained controversial in the deep sea where light is limited. Striatobalanus tenuis (Hoek, 1883) is a deep sea (300 - 400m) acorn barnacle found in East and South China Sea. It inhabits on rocks, gastropod shells and crab surface and is commonly found as a bycatch of bottom trawl in Taiwan. In this study, we collected adult S. tenuis individuals from Kezailiao fishing port and cultivated their fertilized egg masses at 20°C. The hatched larvae were fed with microalgae and water is changed once a week. Precipitation of microalgae, which may disturb larval movement, was frequently removed. Six stages (NI-NVI) of swimming nauplius larvae and one stage of non-swimming cyprid larvae were successfully observed. Different nauplius stages of S. tenuis show morphological features corresponding to "normal" developmental sequence of barnacles. Red, green, blue and white light were applied to test phototaxis of S. tenuis nauplii. NII and NIII larvae were attracted by all the light settings. NIV and NV larvae showed positive phototaxis to all the light settings except blue light, which was negative. NVI larvae were repelled by all the light settings. The change of light preference at different nauplius stages might relate to their sequence of vertical migration. We suggested that the relaeased larvae migrate to shallower water, about 100m deep, to feed phytoplankton, then migrate back to the deep water for settlement.

論文審定書 i
誌謝 ii
摘要 iii
Abstract iv
目錄 v
圖目錄 vii
表目錄 ix
壹、 前言 1
貳、 材料方法 4
一、 藤壺採集及幼生飼育 4
二、 幼生體型觀察及樣本保存 4
三、 幼生附肢剛毛形態觀察－解剖及光學顯微鏡 5
四、 幼生形態觀察－掃瞄式電子顯微鏡 (scanning electron microscope ,SEM) 6
臨界點乾燥 6
覆膜 6
掃瞄式電子顯微鏡 (SEM) 6
五、 趨光性實驗 7
參、 結果 9
一、 幼生飼育及觀察 9
幼生飼育 9
幼生形態描述 9
各時期體型大小之比較 16
二、 趨光性 16
實驗裝置照度測量 16
實驗裝置光譜組成 16
趨光性實驗 17
肆、 討論 23
伍、 參考文獻 31
附錄一、幼生部位中英文對照表 I
附錄二、2017年11月 - 2018年12月薄殼條藤壺採樣紀錄 II
附錄三、宿主清單 III

Anil, A. C., et al. (2001). "Larval development and metamorphosis in Balanus amphitrite Darwin (Cirripedia; Thoracica): significance of food concentration, temperature and nucleic acids." Journal of Experimental Marine Biology and Ecology 263 (2): 125-141.

Barnes, H. and Barnes, M. (1959). "The naupliar stages of Balanus hesperius Pilsbry." Canadian Journal of Zoology 37 (3): 237-244.

Barnes, H. and Crisp, D. (1956). "Evidence of self-fertilization in certain species of barnacles." Journal of the Marine Biological Association of the United Kingdom 35 (3): 631-639.

Bingham, B. L. and Young, C. M. (1993). "Larval phototaxis in barnacles and snails associated with bathyal sea urchins." Deep Sea Research Part I: Oceanographic Research Papers 40 (1): 1-12.

Bouchet, P. and Waren, A. (1979). "Planktotrophig larval development in deep-water gastropods." Sarsia 64 (1-2): 37-40.

Brickner, I. and Høeg , J. T. (2010). "Antennular specialization in cyprids of coral-associated barnacles." Journal of Experimental Marine Biology and Ecology 392 (1-2): 115-124.

Buchanan, C., et al. (1982). "A laboratory method for studying zooplankton swimming behaviors." Hydrobiologia 94 (1): 77-89.

Buhl-Mortensen, L. and Høeg, J. T. (2006). "Reproduction and larval development in three scalpellid barnacles, Scalpellum scalpellum (Linnaeus 1767), Ornatoscalpellum stroemii (M. Sars 1859) and Arcoscalpellum michelottianum (Seguenza 1876), (Crustacea: Cirripedia: Thoracica): implications for reproduction and dispersal in the deep sea." Marine biology 149 (4): 829-844.

Bulkowski, L. and Meade, J. W. (1983). "Changes in phototaxis during early development of walleye." Transactions of the American Fisheries Society 112 (3): 445-447.

Chan, B. K., et al. (2009). Crustacean fauna of Taiwan: barnacles, volume I-Cirripedia: Thoracica excluding the Pyrgomatidae and Acastinae, National Taiwan Ocean University.

Chan, B. K., et al. (2017). "Cirripede Cypris Antennules: How Much Structural Variation Exists Among Balanomorphan Species from Hard-Bottom Habitats?" The Biological Bulletin 233 (2): 135-143.

Chiang, W.-L., et al. (2007). "Are barnacle larvae able to escape from the threat of UV?" Marine biology 151 (2): 703-711.

Chiang, W., et al. (2003). "UV-B damages eyes of barnacle larvae and impairs their photoresponses and settlement success." Environmental science & technology 37 (6): 1089-1092.

Cohen, J. H. and Forward Jr, R. B. (2002). "Spectral sensitivity of vertically migrating marine copepods." The Biological Bulletin 203 (3): 307-314.

Colon-urban, R., et al. (1979). "Observations on the development and maintenance of the deep sea barnacle, Octolasmis aymonini geryonophila (Pilsbry)." International Journal of Invertebrate Reproduction 1 (4): 245-252.

Crippen, R. W. and Perrier, J. (1974). "The use of neutral red and Evans blue for live-dead determinations of marine plankton (with comments on the use of rotenone for inhibition of grazing)." Stain Technology 49 (2): 97-104.

Crisp, D. (1962). "The larval stages of Balanus hameri (Ascanius, 1767)." Crustaceana 4 (2): 123-130.

Dayton, P. K., et al. (1982). "The vertical zonation of the deep-sea Antarctic acorn barnacle, Bathylasma corolliforme (Hoek): experimental transplants from the shelf into shallow water." Journal of Biogeography: 95-109.

Egan, E. and Anderson, D. (1985). "Larval development of Elminius covertus Foster and Hexaminius popeiana Foster (Cirripedia: Archaeobalanidae: Elminiinae) reared in the laboratory." Marine and Freshwater Research 36 (3): 383-404.

Farrapeira, C. M. R. (2010). "Shallow water Cirripedia of the northeastern coast of Brazil: the impact of life history and invasion on biogeography." Journal of Experimental Marine Biology and Ecology 392 (1-2): 210-219.

Forward, R. B. (1990). "Responses of crustacean larvae to hydrostatic pressure: behavioral basis of high barokinesis." Marine & Freshwater Behaviour & Phy 17 (4): 223-232.

Gaonkar, C. A. and Anil, A. C. (2010). "What do barnacle larvae feed on? Implications in biofouling ecology." Journal of the Marine Biological Association of the United Kingdom 90 (6): 1241-1247.

Gehrke, P. (1994). "Influence of light intensity and wavelength on phototactic behaviour of larval silver perch Bidyanus bidyanus and golden perch Macquana ambigua and the effectiveness of light traps." Journal of Fish Biology 44 (5): 741-751.

Glenner, H. and Høeg, J. T. (1993). "Scanning electron microscopy of metamorphosis in four species of barnacles (Cirripedia Thoracica Balanomorpha)." Marine biology 117 (3): 431-439.

Høeg, J. T. (1995). "The biology and life cycle of the Rhizocephala (Cirripedia)." Journal of the Marine Biological Association of the United Kingdom 75 (3): 517-550.

Haeckel, E. (1887). Report on the scientific results of the voyage of HMS Challenger during the years 1873-76: under the command of Captain George S. Nares, RN, FRS and Captain Frank Tourle Thomson, RN. Zoology: 18. Report on the Radiolaria collected by HMS Challenger during the years 1873-1876/by Ernst Haeckel: 2. Osculosa (Nassellaria and Phæodaria), Eyre & Spottiswoode.

Hamasaki, K., et al. (2013). "Ontogeny of sinking velocity, body density, and phototactic behaviour in larvae of the coconut crab Birgus latro: implications for larval dispersal and recruitment in the sea." Journal of Experimental Marine Biology and Ecology 442: 58-65.

Hills, J., et al. (1998). Do barnacle larvae respond to multiple settlement cues over a range of spatial scales? Recruitment, Colonization and Physical-Chemical Forcing in Marine Biological Systems, Springer: 101-111.

Häder, D.-P. and Häder, M. A. (1988). "Inhibition of motility and phototaxis in the green flagellate, Euglena gracilis, by UV-B radiation." Archives of microbiology 150 (1): 20-25.

Jékely, G., et al. (2008). "Mechanism of phototaxis in marine zooplankton." Nature 456 (7220): 395.

Jablonski, D. and Lutz, R. A. (1983). "Larval ecology of marine benthic invertebrates: paleobiological implications." Biological Reviews 58 (1): 21-89.

Jensen, P. G., et al. (1994). "Scanning electron microscopy of lattice organs in cyprids of the Rhizocephala Akentrogonida (Crustacea: Cirripedia)." Canadian Journal of Zoology 72 (6): 1018-1026.

Jensen, P. G., et al. (1994). "Comparative SEM studies of lattice organs: putative sensory structures on the carapace of larvae from Ascothoracida and Cirripedia (Crustacea Maxillopoda Thecostraca)." Acta Zoologica 75 (2): 125-142.

Jones, D. S. and Hosie, A. M. (2016). "A checklist of the barnacles (Cirripedia: Thoracica) of Singapore and neighbouring waters." Raffles Bulletin of Zoology.

Knight-Jones, E. and Morgan, E. (1966). "Responses of marine animals to changes in hydrostatic pressure." Oceanogr. Mar. Biol. Ann. Rev 4: 267-299.

Korn, O. and Rybakov, A. (2001). "Larval development in the rhizocephalan barnacle Sacculina pilosella." Russian Journal of Marine Biology 27 (3): 177-179.

Korn, O. M. and Elfimov, A. S. (1999). "Larval development of a warm-water immigrant barnacle, Solidobalanus fallax (Cirripedia: Archaeobalanidae) reared in the laboratory." Journal of the Marine Biological Association of the United Kingdom 79 (6): 1039-1044.

Lang, W. H. (1979). "Larval development of shallow water barnacles of the Carolinas (Cirripedia: Thoracica) with keys to naupliar stages."

Lang, W. H., et al. (1979). "Behavioral responses of Balanus improvisus nauplii to light intensity and spectrum." The Biological Bulletin 157 (1): 166-181.

Le Tourneux, F. and Bourget, E. (1988). "Importance of physical and biological settlement cues used at different spatial scales by the larvae of Semibalanus balanoides." Marine biology 97 (1): 57-66.

Lee, C. and Shim, J. M. (2000). "Larval development of Membranobalanus koreanus Kim & Kim, 1983, reared in the laboratory (Cirripedia, Thoracica)." Crustaceana-International Journal of Crystacean Research- 73 (4): 393-406.

Lin, H.-C., et al. (2015). "The origins and evolution of dwarf males and habitat use in thoracican barnacles." Molecular phylogenetics and evolution 91: 1-11.

Lin, H., et al. (2017). "A peculiar lens-shaped structure observed in the South China Sea." Scientific reports 7 (1): 478.

Lipps, J. (1982). "Origin, age, and evolution of Antarctic and deep-sea faunas." The Environments of the Deep Sea 2: 325-356.

Marsden, J. R. (1994). "Vertical movements and distribution of planktonic larvae of the serpulid polychaete Spirobranchus polycerus (Schmarda); effects of changes in hydrostatic pressure." Journal of Experimental Marine Biology and Ecology 176 (1): 87-105.

Martin, J. W., et al. (2014). Atlas of crustacean larvae, JHU Press.

Moyse, J. (1961). The larval stages of Acasta spongites and Pyrgoma anglicum (Cirripedia). Proceedings of the Zoological Society of London, Wiley Online Library.

Nasrolahi, A., et al. (2007). "Effects of algal diet on larval survival and growth of the barnacle Amphibalanus (= Balanus) improvisus." Journal of the Marine Biological Association of the United Kingdom 87 (5): 1227-1233.

Newman, W. A. and Ross, A. (2001). "Prospectus on larval cirriped setation formulae, revisited." Journal of Crustacean Biology 21 (1): 56-77.

Obukhova, A., et al. (2015). "The morphology of the frontal filaments in the nauplii of Verruca stroemia (Müller, 1776) and Hesperibalanus hesperius (Pilsbry, 1916) (Cirripedia: Thoracica)." Russian Journal of Marine Biology 41 (5): 363-371.

Pawlik, J. R. (1992). "Chemical ecology of the settlement of benthic marine invertebrates." Oceanography and marine biology: an annual review.

Pfeiffer-Herbert, A. S., et al. (2007). "Dispersal of barnacle larvae along the central California coast: a modeling study." Limnology and oceanography 52 (4): 1559-1569.

Pilsbry, H. A. (1907). The barnacles (Cirripedia) contained in the collections of the US National Museum, US Government Printing Office.

Rainbow, P. and Walker, G. (1976). "The feeding apparatus of the barnacle nauplius larva: a scanning electron microscope study." Journal of the Marine Biological Association of the United Kingdom 56 (2): 321-326.

Rex, M. A. and Warén, A. (1982). "Planktotrophic development in deep-sea prosobranch snails from the western North Atlantic." Deep Sea Research Part A. Oceanographic Research Papers 29 (2): 171-184.

Rumrill, S. S. (1990). "Natural mortality of marine invertebrate larvae." Ophelia 32 (1-2): 163-198.

Rybakov, A. V., et al. (2002). "Larval development in Peltogasterella studied by scanning electron microscopy (Crustacea: Cirripedia: Rhizocephala)." Zoologischer Anzeiger-A Journal of Comparative Zoology 241 (3): 199-221.

Shanks, A. (2001). An identification guide to the larval marine invertebrates of the Pacific Northwest, Oregon State University Press.

Shirley, S. M. and Shirley, T. C. (1988). "Behavior of red king crab larvae: phototaxis, geotaxis and rheotaxis." Marine & Freshwater Behaviour & Phy 13 (4): 369-388.

Singarajah, K., et al. (1967). "The effect of feeding upon the phototactic behaviour of cirripede nauplii." Journal of Experimental Marine Biology and Ecology 1 (2): 144-153.

Sneed, J. M., et al. (2014). "The chemical cue tetrabromopyrrole from a biofilm bacterium induces settlement of multiple Caribbean corals." Proceedings of the Royal Society B: Biological Sciences 281 (1786): 20133086.

Southward, A. J. (1975). "Intertidal and shallow water Cirripedia of the Caribbean." Studies on the fauna of Curaçao and other Caribbean Islands 46 (1): 1-53.

Storz, U. and Paul, R. (1998). "Phototaxis in water fleas (Daphnia magna) is differently influenced by visible and UV light." Journal of Comparative Physiology A 183 (6): 709-717.

Thiyagarajan, V., et al. (1997). "Larval description of Balanus reticulatus Utinomi (Cirripedia, Balanidae), reared in the laboratory." Journal of Experimental Marine Biology and Ecology 209 (1-2): 215-231.

Thorson, G. (1936). The larval development, growth, and metabolism of arctic marine bottom invertebrates compared with those of other seas, CA Reitzel.

Thorson, G. (1964). "Light as an ecological factor in the dispersal and settlement of larvae of marine bottom invertebrates." Ophelia 1 (1): 167-208.

Walker, G. (1973). "Frontal horns and associated gland cells of the nauplii of the barnacles, Balanus hameri, Balanus balanoides and Elminius modestus (Crustacea: Cirripedia)." Journal of the Marine Biological Association of the United Kingdom 53 (2): 455-463.

Watanabe, H., et al. (2004). "Larval development and intermoult period of the hydrothermal vent barnacle Neoverruca sp." Journal of the Marine Biological Association of the United Kingdom 84 (4): 743-745.

Wong, G. T., et al. (2015). "Hydrography and nutrient dynamics in the northern South China sea Shelf-sea (NoSoCS)." Deep Sea Research Part II: Topical Studies in Oceanography 117: 23-40.

Wu, R., et al. (1997). "A phototaxis inhibition assay using barnacle larvae." Environmental Toxicology and Water Quality: An International Journal 12 (3): 231-236.

Yan, Y. and Chan, B. K. (2001). "Larval development of Chthamalus malayensis (Cirripedia: Thoracica) reared in the laboratory." Journal of the Marine Biological Association of the United Kingdom 81 (4): 623-632.

Yap, F., et al. (2015). "Larval development of the pedunculate barnacles Octolasmis angulata Aurivillius 1894 and Octolasmis cor Aurivillius 1892 (Cirripedia: Thoracica: Poecilasmatidae) from the gills of the mud crab, Scylla tranquebarica Fabricius, 1798." Arthropod structure & development 44 (3): 253-279.

Zardus, J. D. and Hadfield, M. G. (2004). "Larval development and complemental males in Chelonibia testudinaria, a barnacle commensal with sea turtles." Journal of Crustacean Biology 24 (3): 409-421.

Zhu, J., et al. (2013). "On summer stratification and tidal mixing in the Taiwan Strait." Frontiers of Earth Science 7 (2): 141-150.

劉瑞玉、任先秋(2007)。 中國動物誌-無脊椎動物第四十二卷 (甲殼動物亞門, 蔓足下綱, 圍胸總目), 北京: 科學出版社.

趙文榮、曾金成、陶申秋 (2011)。 餌料生物學-角毛藻之培養 I:43-46, 儒林出版社。台北。

自然與人文數位博物館，台灣。 Retrieved November 09, 2017, from http://digimuse.nmns.edu.tw/Demo_2011/NewModule.aspx?ObjectId=
0b00000181a9f4c5&ParentID=0b00000181a9f4c5&Type=&Part=&Domain=az&Field=ar&Language=CHI

周芷儀、李念宜、陳紫媖 (2017)。 牟氏角毛藻及擬球藻之培養 行政院農委會水產實驗所電子報第138期 Retrieved July 12, 2019, from http://
www.tfrin.gov.tw/friweb/frienews/enews0138/c5.html

臺灣生物多樣性資訊網，台灣。 Retrieved J 09, 2018, from http://taibif.tw/

科技部海洋科學學門資料庫，台灣。 Retrieved November 09, 2017, from http://www.odb.ntu.edu.tw/ctd/display/ctd15moa/