石首魚（Sciaenidae）雄魚在繁殖期時會藉由發音肌振動，並同時利用泳鰾共鳴發出聲響；此外在受到驚擾及攝食時亦會發出聲響。大黃魚（Larimichthys crocea）是石首魚類，主要分佈於南中國海、東海及黃海南部一帶，為中國沿海極為重要的經濟魚種之一；過去由於其重要的經濟效應而被過度捕撈以至於在野外數量大減。近年來人工繁殖技術成熟，養殖場對重要的經濟石首魚類進行養殖，大黃魚亦在中國大陸福建省被成功的人工繁殖並大量飼養。由於大黃魚為石首魚科中少數雌、雄魚皆具發音肌的魚種，牠提供探討不同性別在發聲特徵上差異的研究材料。本研究針對大黃魚在受到人為干擾時所發出之聲音，及繁殖期（求偶、產卵、產卵後）的聲訊變化做描述，同時藉由分子親緣關係鑑定了解大黃魚在石首魚科內所處的演化地位。並探討石首魚發音構造之形態（泳鰾形狀）的變異是否與其演化途徑相符。結果發現，大黃魚之干擾聲音在雌、雄魚間有顯著的差異。此外生殖聲音及干擾聲音在特徵上亦有明顯的差別。大黃魚受到驚擾時所發出的聲音其脈衝數最多可高達二十三個。以經注射激素（LHRH-A3）之標本進行實驗得到結果顯示，在僅單有雄性或雌性的池子中所錄到的生殖聲音為僅含一到兩個脈衝數的聲音，而在雌雄魚混養的池中，其生殖聲音約從下午兩時至三時左右開始出現，每一個叫聲是由一到七個脈衝所組成；含有越多脈衝的聲音出現機率隨著夜間的時序而增加，於生殖聲音剛開始出現時，僅一個脈衝的聲音出現機率相當高，在高達七個脈衝的聲音出現之後即可在養殖池中觀察到大量卵的出現。推測大黃魚的生殖聲音有促進母魚排卵的功能。親緣關係分析結果顯示，梅童魚（Collichthys）為大黃魚之姊妹群，其泳鰾形態與大黃魚相同（邊緣有許多樹枝狀附枝）。 此外，石首魚類泳鰾之型態由簡單不具附枝演化成有許多附枝的複雜型態。Larimichthys 和 Collichthys，與其它分佈於印度西太平洋的石首魚屬，包括Miichthys、Boesemania、Bahaba、Panna、Atrobucca、Otolithes、Pterotolithus、Chrysochir、Paranebris、Protonibea、Pennahia、Nibea、Dendrophysa、Johnius等具近親緣關係而組成一單系群（bootstrap value=100）。其中除Boesemania、Bahaba及Paranebris的泳鰾較為簡單外，泳鰾周圍皆具許多附枝。由此推測此三屬的泳鰾可能因逆轉而產生。

The fishes of the family Sciaenidae have been known to vocalize during the reproductive season, and it is known that in most species only the male calls. Sounds are produced by vibration of the sonic muscles, which set the swim bladder into resonance. In addition, they also emit sounds by the same mechanism while being disturbed. The large yellow croaker, Larimichthys crocea (Perciformes, Sciaenidae) is one of the important commercial fish species distributed in South China Sea, East China Sea and southern Yellow Sea. In the past years, they have been overfished because of their high fishery value. Many aquatic farms started to culture economically important sciaenids because of the established artificial propagation technique. Now large yellow croakers have been successfully cultured in Fuchien Province, China. Unlike most sciaenids, the sonic muscles are only possessed by male, both male and female large yellow croakers have sonic muscles. This species provides the best opportunity to investigate the characteristics of sounds produced by different genders. The aims of this study were 1) to describe the hand-held disturbance sounds in large yellow croakers, 2) to describe the sounds produced during courtship and spawning in large yellow croakers being injected hormone (LHRH-A3), 3) to investigate the phylogenetic position of large yellow croaker in relation to other sciaenid fishes, and 4) to understand the evolutionary path of swim bladder morphology in the family Sciaenidae. The results show that 1) the pulse numbers of hand-held disturbance sounds in large yellow croaker can reach 23; 2) The reproductive sounds consisted of 1 to 7 pulses which started at about 1400 hr, and both vocal activity (no. sounds/min) and pulse numbers per call would increase with time. However, spawning occupied at the time slot when pulse numbers per call reached 7 in the unisex pond. The sounds in the ponds with only males or females can only recorded the sounds with 1 to 2 pulses; 3) The genus Collichthys was the sister taxon of large yellow croaker. Morphology of the swim bladder in Collichthys is similar to large yellow croaker; 4) Morphology of swim bladder evolved from simple to complex forms. Finally, Larimichthys, Collichthys, and other sciaenid genera distributed in the Indo-western Pacific Ocean including Miichthys, Boesemania, Bahaba, Panna, Atrobucca, Otolithes, Pterotolithus, Chrysochir, Paranebris, Protonibea, Pennahia, Nibea, Dendrophysa and Johnius form a monophyletic group with a bootstrap value of 100. Most of the members have the complex swim bladders with many appendages except Boesemania, Bahaba and Paranebris. Presence of simple form of the swim bladder in these three genera is inferred as a result of morphological reversal.

Introduction 1
Material and methods 6
2.1 Study species 6
2.2 Recording and analysis of sounds 7
2.2.1 Audio recording equipment 7
2.2.2 Acoustic analyses 8
2.2.3 Observation on spawning 8
2.2.4 Statistical analyses 9
2.3 Phylogenetic analysis 9
2.3.1 Taxonomic sampling 10
2.3.2 DNA extraction 10
2.3.3 PCR condition 11
2.4 Morphology comparison 12
Results 13
3.1 Acoustic signals 13
3.1.1. Hand-held disturbance sounds 13
3.1.1.1. Characteristics 13
3.1.1.2. Difference between the hand-held disturbance sounds of two genders 14
3.1.2. Reproductive sounds 15
3.1.2.1. Characteristics 15
3.1.2.2. Temporal change of sound types in an evening when spawning took place 16
3.1.3. Difference between disturbance and reproductive sounds 17
3.2 Phylogenetic relationships 17
Discussion 18
4.1 Acoustic signals 18
4.1.1 Variation in sound characteristics 18
4.1.2 Sound patterns during reproduction 19
4.1.3 Function of reproductive sounds 19
4.2 Evolutionary significance of swim bladder morphology 20
4.2.1 Phylogeny 20
4.2.2 Implications for acoustic evolution based on swim bladder morphology 21
Conclusion 22
References 24

Aalbers, S. A. 2008. Seasonal, diel, and lunar spawning periodicities and associated sound production of white seabass (Atractoscion nobilis). Fishery Bulletin 106:143-151.
Aalbers, S. A., and M. A. Drawbridge. 2008. White seabass spawning behavior and sound production. Transactions of the American Fisheries Society 137:542-550.
Amorim, M. 2000. Growling for food: acoustic emissions during competitive feeding of the streaked gurnard. Journal of Fish Biology 57:895-907.
Avise, J. C., and D. Ellis. 1986. Mitochondrial DNA and the evolutionary genetics of higher animals Philosophical Transactions of the Royal Society B: Biological Sciences 312:325-342.
Barimo, J. F., and M. L. Fine. 1998. Relationship of swim-bladder shape to the directionality pattern of underwater sound in the oyster toadfish. Canadian Journal of Zoology 134:134-143.
Burkenroad, M. D. 1931. Notes on the sound-producing marine fishes of Louisiana. Copeia 1931:20-27.
Chao, L. N. 1978. A Basis For Classifying Western Atlantic Sciaenidae (Teleostei: Perciformes). National Marine Fisheries Service, National Oceanic and Atmospheric Administration, United States Dept. of Commerce.
Chao, L. N. 1986. A synopsis on zoogeography of the Sciaenidae. Pages 570-589 In T. Uyeno, R. Arai, T. Taniuchi, and K. Matsuura (Eds.). Indo-Pacific Fish Biology: Proceedings of the Second International Conference on Indo-Pacific fishes. Ichthyological Society of Japan. Tokyo.
Chen, W.-J., C. Bonillo, and G. Lecointre. 2003. Repeatability of clades as a criterion of reliability: a case study for molecular phylogeny of Acanthomorpha (Teleostei) with larger number of taxa. Molecular Phylogenetics and Evolution 26:262-288.
Chen, W., M. Miya, K. Saitoh, and R. Mayden. 2008. Phylogenetic utility of two existing and four novel nuclear gene loci in reconstructing tree of life of ray-finned fishes: the order Cypriniformes (Ostariophysi) as a case study. Gene 423:125-134.
Chen, W. J., R. Ruiz-Carus, and G. Orti. 2007. Relationships among four genera of mojarras (Teleostei: Perciformes: Gerreidae) from the western Atlantic and their tentative placement among percomorph fishes. Journal of Fish Biology 70:202-218.
Cheng, Y., T. Xu, G. Shi, and R. Wang. 2010. Complete mitochondrial genome of the miiuy croaker Miichthys miiuy (Perciformes, Sciaenidae) with phylogenetic consideration. Marine Genomics 3:201-209.
Colson, D. J., S. N. Patek, E. L. Brainerd, and S. M. Lewis. 1998. Sound production during feeding in Hippocampus seahorses (Syngnathidae). Environmental Biology of Fishes 51:221-229.
Connaughton, M. A., and M. H. Taylor. 1994. Seasonal cycles in the sonic muscles of the weakfish, Cynosion regalis. Fishery Bulletin 92:697-703.
Connaughton, M. A., and M. H. Taylor. 1995. Seasonal and daily cycles in sound production associated with spawningin the weakfish, Cynoscion regalis. Environmental Biology of Fishes 42:233-240.
Connaughton, M. A., and M. H. Taylor. 1996. Drumming courtship, and spawning behavior in captice weakfish, Cynoscion regalis. Copeia 1996:195-199.
Connaughton, M. A., M. H. Taylor, and M. L. Fine. 2000. Effects of fish size and temperature on weakfish disturbance calls: implications for the mechanism of sound generation. The Journal of Experimental Biology 203:2503-2512.
Connaughton, M. A., M. L. Fine, and M. H. Taylor. 2002. Weakfish sonic muscle: influence of size, temperature and season. The Journal of Experimental Biology 205:2183-2188.
Connaughton, M. A. 2004. Sound generation in the searobin (Prionotus carolinus), a fish with alternate sonic muscle contraction. Journal of Experimental Biology 207:1643-1654.
Cui, Z., Y. Liu, C. Li, F. You, and K. Chu. 2009. The complete mitochondrial genome of the large yellow croaker, Larimichthys crocea (Perciformes, Sciaenidae): unusual features of its control region and the phylogenetic position of the Sciaenidae. Gene 432:33-43.
Demski, L. S., J. W. Gerald, and A. N. Popper. 1973. Central and peripheral mechanisms of teleost sound production. American Zoologist 13:1141-1168.
Felsenstein, J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39:783-791.
Felsentein, J. 1981. Evolutionary trees from DNA sequences: a maximum likelihood approach. Journal of Molecular Evolution 17:368-376.
Feng, Z., and Q. Cao. 1979. Ichthyology. Agricultural Press House, Beijing.
Fine, M. L., D. McElroy, J. Rafi, C. B. King, K. E. Losser, and S. Newton. 1996. lateralization of pectoral stridulation sound production in the channel catfish. Physiology and Behavior 60:753-757.
Fish, J. F., and W. C. Cummings. 1972. A 50-dB increase in sustained ambient noise from fish Cynoscion xanthulus. Journal of Acoustical Society of America 52:1266-1269.
Fish, M. P., and W. H. Mowbray. 1970. Sounds of Western North Atlantic fishes. Johns Hopkins Press, Baltimore.
Gainer, H. 1967. Neuromuscular mechanisms of sound production and pectoral spine locking in the banjo catfish, Bunocephalus species. Physiological Zoology 40:296-306.
Gannon, D. P. 2007. Acoustic behavior of Atlantic croaker, Micropogonias undulatus (Sciaenidae). Copeia 2007:193-204.
Hawkins, A. D., and M. C. P. Amorima. 2000. Spawning sounds of the male haddock, Melanogrammus aeglefinus. Environmental Biology of Fishes 59:30-42.
Hill, G. L., M. L. Fine, and J. A. Musick. 1987. Ontogeny of the sexually dimorphic sonic muscle in three Sciaenid species. Copeia 1987:708-713.
Kaatz, I. M. 2002. Multiple sound-producing mechanisms in teleost fishes and hypotheses regarding their behavioural significance. Bioacoustics 12:230-233.
Kasumyan, A. O. 2009. Acoustic signaling in fish. Journal of Ichthyology 49:963-1020.
Lopez, J. A., W.-J. Chen, and G. Orti. 2004. Esociform phylogeny. Copeia 2004:449-464.
Ladich, F. 2004. Sound production and acoustic communication. Pages 210-230 In G. V. D. Emde, J. Mogdans, and B. G. Kapoor (Eds.). The senses of fish: adaptations for the reception of natural stimuli. Narosa, New Delhi.
Lagardere, J. P., and A. Mariani. 2006. Spawning sounds in meagre Argyrosomus regius recorded in the Gironde estuary, France. Journal of Fish Biology 69:1697-1708.
Lin, Y. C., H. K. Mok, and B. Q. Huang. 2007. Sound characteristics of big-snout croaker, Johnius macrorhynus (Sciaenidae). The Journal of the Acoustical Society of America 121:586-593.
Lobel, P. S. 1992. Sounds produced by spawning fishes. Environmental Biology of Fishes 33:351-358.
Lobel, P. S. 2002. Diversity of fish spawning sounds and the application of passive acoustic monitoring. Bioacoustics 12:286-289.
Luczkovich, J. J., M. W. Sprague, S. E. Johnson, and R. C. Pullinger. 1999. Delimiting spawning areas of weakfish, Cynoscion regalis (family Sciaenidae) in Pamlico Sound, North Carolina using passive hydroacoustic surveys. Bioacoustics 10:143-160.
Mohan, R. S. L. 1972. A synopsis of the Indian genera of the fishes of the Family Sciaenidae. Indian Journal of Fisheries 16:82-98.
Mok, H. K., and R. G. Gilmore. 1983. Analysis of sound production in estuarine aggregations of Pogonias cromis, Bairdiella chrysoura, and Cynoscion nebulosus (Sciaenidae). Bulletin of the Institute of Zoology, Academia Sinica 22:157-185.
Myrberg, A. 1997. Underwater sound: Its relevance to behavioral functions among fishes and marine mammals. Marine and Freshwater Behaviour and Physiology 29:3-21.
Parmentier, E. 2006. Sound production mechanism in carapid fish: first example with a slow sonic muscle. Journal of Experimental Biology 209:2952-2960.
Parmentier, E., P. Vandewalle, C. Brie, L. Dinraths, and D. Lecchini. 2010. Comparative study on sound production in different Holocentridae species. Frontiers in Zoology 8:1-12.
Popper, A. N., R. R. Fay, C. Platt, and O. Sand. 2003. Sound detection mechanisms and capabilities of teleost fishes. Pages 3-38 In S. P. Collin, and N. J. Marshall (Eds.). Sensory processing in aquatic environments. Springer-Verlag New York, Inc., New York.
Rambaut, A. 1996. Sequence Alignment Editor. Department of Zoology University of Oxford, U.K.
Ramcharitara, J., D. P. Gannonb, and A. N. Popper. 2006. Bioacoustics of fishes of the Family Sciaenidae (croakers and drums). Transactions of the American Fisheries Society 135:1409-1431.
Ren, X. M., D. Z. Gao, Y. L. Yao, F. Yang, J. F. Liu, and F. J. Xie. 2007. Occurrence and characteristic of sound in large yellow croaker (Pseudosciaena crocea). Journal oF Dalian Fisheries University 22:123-128.
Richardson, J. 1846. Report on the ichthyology of the Seas of China and Japan. Report of the British Association for the Advancement of Science 1845:187-320.
Ripley, J. L., P. S. Lobel, and H. Y. Yan. 2002. Coorelation of sound production with hearing sesnsitivity in the Lake Malawi cichlid Tramitichromis intermedius. Bioacoustics 12:238-240.
Sasaki, K. 1989. Phylogeny of the Family Sciaenidae, with notes on its zoogeography (Teleostei, Perciformes). Pages 1-137 in. Memoirs of the Faculty of Fisheries.
Saucier, M. H., and D. M. Baltz. 1993. Spawning site selection by spotted seatrout, Cynoscion nebulosus, and black drum, Pogonias cromis, in Louisiana. Environmental Biology of Fishes 36:257-262.
Schwarzhans, W. 1993. A comparative morphological treatise of recent and fossil otoliths of the Family Sciaenidae (Perciformes) (Piscium catalogus), Vol. 1. Verlag F. Pfeil Munche.
Sebeok, T. A. 1977. How Animals Communicate. Indiana University Press, Bloomington.
Stamatakis, A. 2006. RAxML-VI-HPC: maximum likelihood-based phylogentic analyses with thousands of taxa and mixed models. Bioinformatics 22:2688-2690.
Stamatakis, A., P. Hoover, and J. Rougemont. 2008. A rapid bootstrap algorithm for the RAxML web-servers. Systematic Biology 5:758-771.
Takemura, A., T. Takita, and K. Mizue. 1978. Underwater calls of the Japanese marine drum fishes (Sciaenidae). Bulletin of the Japanese Society of Scientific Fisheries 44:121-125.
Tellechea, J. S., C. Martinez, M. L. Fine, and W. Norbis. 2010. Sound production in the whitemouth croaker and relationship between fish size and disturbance call characteristics. Environmental Biology of Fishes 89:163-172.
Tellechea, J. S., W. Norbis, D. Olsson, and M. L. Fine. 2011. Calls of the black drum (Pogonias cromis: Sciaenidae): geographical differences in sound production between Northern and Southern hemisphere populations. Journal of Experimental Zoology 315:48-55.
Tower, R. W. 1908. The production of sound in the drumfishes, the sea-robin and the toadfish. Annals of the New York Academy of Sciences 18:149-180.
Trewavas, E. 1977. The sciaenid fishes (croakers or drums) of the Indo-West-Pacific. The Transaction of the Zoological Society of Lodon 33:253-541.
Ueng, J. P., B. Q. Huang, and H. K. Mok. 2007. Sexual differences in the spawning sounds of the Japanese croaker, Argyrosomus japonicus (Sciaenidae). Zoological Studies 46:103-110.
Vergara-Chen, C., W. E. Aguirre, M. Gonzalez-Wanguemert, and E. Bermingham. 2009. A mitochondrial DNA based phylogeny of weakfish species of the Cynoscion group (Pisces: Sciaenidae). Molecular Phylogenetics and Evolution 53:602-607.
Wang, Z., Y. Wang, L. Lin, S. Khoo, and H. Okamoto. 2002. Genetic polymorphism in wild and cultured large yellow croaker Pseudosciaena crocea using AFLP fingerprinting. Journal of Fishery Sciences of China 9:199-203.
Ward, R. D., T. S. Zemlak, B. H. Innes, P. R. Last, and P. D. N. Hebert. 2005. DNA barcoding Australia's fish species. Philosophical Transactions of the Royal Society B: Biological Sciences 360:1847-1857.
Wilson, B., R. S. Batty, and L. M. Dill. 2004. Pacific and Atlantic herring produce burst pulse sounds. Proceedings of the Royal Society B: Biological Sciences 271:95-97.
Yan, H. Y., M. L. Fine, N. S. Horn, and W. E. Colon. 2000. Variability in the role of the gasbladder in fish audition. Journal of Comparative Physiology A: Sensory, Neural, and Behavioral Physiology 186:435-445.
Zelick, R., D. Mann, and A. N. Popper. 1999. Acoustic communication in fishes and frogs. Pages 363-411 In R. R. Fay, and A. N. Popper (Eds.). Comparative Hearing: Fish and Amphibians, Vol. 11. Springer, New York.
Zhao, S. L., R. X. Wang, and X. S. Liu. 2002. Reasons of exhaustion of resources of Pseudosciaena crocea in Zhoushan fishing ground and the measures of protection and proliferation. Journal of Zhejiang Ocean University 21:160-165.