Bellorin, A.M., M.C. Oliveira and E.C. Oliveira. 2002. Phylogeny and systematics of the marine algal family Gracilariaceae (Gracilariales, Rhodophyta) based on small subunit rDNA and ITS sequences of Atlantic and Pacific species. J. Phycol., 38: 551-563.
Bellorin, A.M., A. Buriyo, J. Sohrabipour, M.C. Oliveira and E.C. Oliveira. 2008.
Gracilariopsis mclachlanii sp. nov. and Gracilariopsis persica sp. nov. of the Gracilariaceae (Gracilariales, Rhodophyceae) from the Indian Ocean. Journal of Phycology, 44: 1022–1032.
Bird, C.J. 1995. A review of recent taxonomic concepts and developments in the Gracilariaceae (Rhodophyta). Journal of Applied Phycology, 7: 255-267.
Byrne, K., G.C. Zuccarello, J. West, M.L. Liao and G.T. Kraft. 2002. Gracilaria species (Gracilariaceae, Rhodophyta) from southeastern Australia, including a new species, Gracilaria perplexa sp. nov.: Morphology, molecular relationships and agar content.
Phycological Research, 50(4): 295-312.
Chang, C.F. and B.M. Xia. 1988. On two new Gracilaria (Gigartinales, Rhodophyta) from South China. In: Taxonomy of economic seaweeds with reference to some Pacific and Caribbean species (Abbott, I.A. Eds) Vol.2, pp. 127-129. California Sea Grant College Program.
Clarkston, B.E. and G.W. Saunders. 2010. A comparison of two DNA barcode markers for species discrimination in the red algal family Kallymeniaceae (Gigartinales,
Florideophyceae), with a description of Euthora timburtonii sp. nov. Botany, 88: 119-131.
- 47 -
Cohen, S., S. Faugeron, E.A. Martines, J.A. Correa, F. Viard, C. Destombe and M. Valero.
2004. Molecular identification of two sibling species under the name Gracilaria chilensis (Rhodophyta, Gracilariales). J. Phycol., 40: 742-747.
Dawson, E.Y. 1949. Studies of northeast Pacific Gracialriaceae. Allan Hancock Foundation Publ., Occ. Pap., 7:1-105, 25pls.
Fredericq, S. and J.N. Norris. 1992. Studies on cylindrical species of western Atlantic Gracilaria (Gracilariales, Rhodophyta): G. cylindrica Børgesen and G. blodgettii Harvey. In: Taxonomy of Economic Seaweeds. (Abbott, I.A. Eds) Vol.2, pp. 211-231.
La Jolla, California: California Sea Grant College.
Fredericq, S. and M.H. Hommersand. 1989. Proposal of the Gracilariales ord. nov.
(Rhodophyta) based on an analysis of the reproductive development of Gracilaria verrucosa. Journal of Phycology, 25: 213-227.
Fredericq, S. and M.H. Hommersand. 1990a. Comparative morphology and taxonomic status of Gracilariopsis (Gracilariales, Rhodophyta). J. Phycol., 25: 228-241.
Fredericq, S. and M.H. Hommersand. 1990b. Diagnoses and key to the genera of the Gracilariaceae (Gracilariales, Rhodophyta). Hydrobiologia, 204/205: 172-178.
Gall, L.L. and G.W. Saunders. 2010. DNA barcoding is a powerful tool to uncover algal diversity: a case study of the Phyllophoraceae (Gigartinales, Rhodophyta) in the Canadian flora. J. Phycol., 46: 374-386.
Gargiulo, G M., M. Morabito, G. Genoves and F.D. Masi. 2006. Molecular systematic and phylogenetics of Gracilariacean species from the Mediterranean Sea. J. Phycol., 18:
297–504.
Gavio, B and S. Fredericq. 2002. Grateloupia turuturu (Halymeniaceae, Rhodophyta) is the correct name of the non-native species in the Atlantic known as Grateloupia doryphora. European Journal of Phycology, 37: 349-360.
- 48 -
Goff, L.J., D.A. Moon, A.W. Coleman. 1994. Molecular delineation of species and species relationships in the red algal agarophytes Gracilariopsis and Gracilaria (Gracilariales). J. phycol., 30: 521-537.
Greville, R.K. 1830. Algae britannicae, or descriptions of the marine and other inarticulated plants of the British islands, belonging to the order Algae; with plates illustrative of the genera. pp. [i*-iii*], [i]-lxxxviii, [1]-218, pl. 1-19.
Guiry, M.D. and G.M. Guiry. 2011. AlgaeBase. World-wide electronic publication, National University of Ireland, Galway. http://www.algaebase.org; searched on 08 December 2011.
Gurgel, C.F.D. and S. Fredericq. 2004. Systematics of the Gracilariaceae (Gracilariales, Rhodophyta): a critical assessment based on rbcL sequence analysis. J. Phycol., 40:
138-159.
Hajibabaei, M., G.A.C. Singer, P.D.N. Hebert and D.A. Hickey. 2007. DNA barcoding:
how it complements taxonomy, molecular phylogenetics and population genetics.
Trends in Genetics, 23: 167–172.
Hall, T.A. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucl Acids Symp Ser, 41: 95-98.
Harvey, W.H. 1855. Short characters of some new genera and species of algae discovered on the coast of the colony of Victoria, Australia. Ann. Mag. Nat. Hist. (Lond.), 5: 332-6.
Hau, L.N. and S.M. Lin. 2006. Gracilariopsis nhatrangensis (Gracilariaceae,
Rhodophyta), a new marine red alga from Nhatrang, southern Vietnam. Botanical studies, 47: 329 –337.
Hebert, P.D N., A. Cywinska, B.L. Ball and J.R. DeWaard. 2003. Biological identification through DNA barcodes. Proc. R. Soc. Lond. B. Biol. Sci., 270(1512): 313-321.
- 49 -
Hebert P.D.N., M.Y. Stoeckle, T.S. Zemlak and C.M. Francis. 2004. Identification of birds through DNA barcodes. PLoS. Biol., 2: 1657-1663.
Iyer, R., E.M. Tronchin, J.J. Bolton and V.E. Coyne. 2005. Molecular systematics of the Gracilariaceae (Gracilariales, Rhodophyta) with emphasis on Southern Africa. J.
Phycol., 41: 672-684.
Kim, M.S, E.C. Yang and S.M. Boo. 2006. Taxonomy and phylogeny of flattened species of Gracilaria (Gracilariceae, Rhodophyta) from Korea based on morphology and protein-coding plastid rbcL and psbA sequences. Phycologia, 45: 520-528.
Kim, M.S., M. Kim, R. Terada, E.C. Yang and S.M. Boo. 2008a. Gracilaria parvispora is the correct name of the species known as G. bursa-pastoris in Korea and Japan. Taxon, 57(1): 231-237.
Kim, M.S., E.C. Yang, S.Y. Kim, I.K. Hwan.g and S.M. Boo. 2008b. Reinstatement of Gracilariopsis chorda (Gracilariaceae, Rhodophyta) based on plastid rbcL and mitochondrial cox1 sequences. Algae, 23: 209-217.
Kim, M.S., M.Y. Yang and G.Y. Cho. 2010. Applying DNA barcoding to Korean Gracilariaceae (Rhdophyta). Cryptogamie, Algologie, 31: 387-401.
Lane, C.E., S.C. Lindstrom and G.W. Saunders. 2007. A molecular assessment of northeast Pacific Alaria species (Laminariales, Phaeophyceae) with reference to the utility of DNA barcoding . Molecular Phylogenetics and Evolution, 44: 634-648.
Lara, A., J.L.P. De Leon, R.Rodriguez, D. Casane, G. Cote, L. Bernatchez and E. Garcia-Machado. 2010. DNA barcoding of Cuban freshwater fishes: evidence for cryptic species and taxonomic conflicts. Molecular Ecology Resources, 10: 421-430.
Lin, S.M. 2008. Morphological and phylogenetic studies of Gracilariopsis chiangii, new species (Gracilariaceae, Rhodophyta), an alga presently known as Gracilaria chorda in Taiwan. The Raffles Bulletin of Zoology Supplement, 19: 19-26.
- 50 -
Lukhtanov, V.A., A. Sourakov, E.V. Zakharov and P.D.N. Hebert. 2009. DNA barcoding central Asian butterflies: increasing geographical dimension does not significantly reduce the success of species identification. Molecular Ecology Resources, 9: 1302-1310.
Marshall, E. 2005. Taxonomy, Will DNA barcodes breathe life into classification?
Science, 307: 1037.
Meier, R., K. Shiyang, G. Vaidya and P.K.L. Ng. 2006. DNA barcoding and taxonomy in Diptera: a tale of high intraspecific variability and low identification success. Syst.
Biol., 55:715-728.
Meyer, C.P. and G. Paulay. 2005. DNA barcoding: error rates based on comprehensive sampling. PLoS. Biol., 3:422.
Millar, A.J.K. and B.M. Xia. 1997. Studies on terete species of Australian Gracilaria. In:
Taxonomy of Economic Seaweeds. (Abbott, I.A. Eds) Vol.6, pp. 103-109. La Jolla, California: California Sea Grant College System.
Montagne, C. 1842. Prodromus generum specierumque phycearum novarum, in itinere ad polum antarcticum... collectarum…. Paris. 16 pp.
Montagne, C. 1843. Quatrième centurie de plantes cellulaires exotiques nouvelles.
Décades VII, IX et X. Ann. Sci. Nat. Bot. Seconde Sér. 20: 352-379, pls 15-16.
Moritz C., C. Cicero. 2004. DNA barcoding: promise and pitfalls. PLoS. Biol., 2:1529-1531.
Nägeli, C. 1847. Die neuern Algensysteme und Versuch zur Begründung eines eigenen Systems der Algen und Florideen. Neue Denkschriften der Allgemeinen
Schweizerischen Gesellschaft für die Gesammten, 9(2): 1-275, pls I-X.
Ohno, M., R. Terada and H. Yamamoto. 1999. The species of Gracilaria from Vietnam.
In: Taxonomy of Economic Seaweeds with reference to some Pacific species, Vol. VII.
- 51 - (Abbott, I.A. Eds) Vol.7, pp. 99-111.
Posada, D. and K.A. Crandall. 1998. MODELTEST: testing the model of DNA substitution. Bioinformatics, 14:817-8.
Robba, L., S.J. Russell, G.L. Barker and J. Brodie. 2006. Assessing the use of the
mitochondrial cox1 marker for use in DNA barcoding of red algae (Rhodophyta). Am.
J. Bot., 93: 1101-1108.
Rueness, J. 2005. Life history and molecular sequences of Gracilaria vermiculophylla (Gracilariales, Rhodophyta), a new introduction to European water. Phycologia., 44:
120-128.
Saunders, G.W. 2005. Applying DNA barcoding to red macroalgae: a preliminary appraisal holds promise for future applications. Philosophical Transactions of the Royal Society of London Series B: Biological Sciences, 360: 1879–1888.
Saunders, G.W. 2008. A DNA barcode examination of the red algal family Dumontiaceae in Canadian waters reveals substantial cryptic species diversity. 1. The foliose Dilsea–
Neodilsea complex and Weeksia. Botany, 86: 773–789.
Sherwood, A.R., M.L. Vis., T.J. Entwisle, O. Necchi and G.G. Presting. 2008. Contrasting intra versus interspecies DNA sequence variation for representatives of the
Batrachospermales (Rhodophyta): Insights from a DNA barcoding approach. Phycol.
Res., 56(4): 269-279.
Sherwood, A.R., A. Kurihara and K.Y. Conklin. 2011. Molecular diversity of Amansieae (Ceramiales, Rhodophyta) from the Hawaiian Islands: a multi-marker assessment reveals high diversity within Amansia glomerata. Phycol. Res., 59: 16.
Swofford D.L. 2002. PAUP*. Phylogenetic Analysis Using Parsimony (*and other Methods). Version 4.0b10. Sinauer Associates, Sunderland, Massachusetts.
Tamura, K., J. Dudley, M. Nei and S. Kumar. 2007. MEGA 4: Molecular Evolutionary
- 52 -
Genetic Analysis (MEGA) Software version 4.0. Mol. Biol. Evol., 24: 1596-1599.
Teasdale, B.W. and A.S. Klein. 2010. Genetic variation and biogeographical boundaries within the red alga Porphyra umbilicalis (Bangiales, Rhodophyta). Bot. Mar., 53(5):
417-431.
Terada, R. and Yamamoto, H. 2000. A taxonomic study on two Japanese species of Gracilaria: Gracilaria shimodensis sp. nov. and Gracilaria blodgettii (Gracilariales, Rhodophyta). Phycol. Res., 48: 189-198.
Terada R., K. Lewmonomont, A. Chirapart and S. Kawaguchi. 2004. Gracilaria and related genera (Gracilariales, Rhodophyta) from the Gulf of Thailand and adjacent waters. Proceeding of the First Joint Seminar on Coastal Oceanography. 144-159.
Terada, R. and S. Shimada. 2005. Taxonomic note on Gracilaria articulata Chang et Xia (Gracilariales, Rhodophyta) from Okinawa, Japan. Cryptogamie Algologie, 26: 77-89.
Walker, R.H., J. Brodie, S. Russell, L.M. Irvine and S. Orfanidis. 2009. Biodiversity of coralline algae in the Northeast Atlantic including Corallina caespitosa sp. nov.
(Corallinoideae, Rhodophyta). J. Phycol., 45(1): 287-297.
Wiemers, M., K. Fiedler. 2007. Does the DNA barcoding gap exist?-a case study in blue butterflies (Lepidoptera: Lycaenidae). Front. Zool., 4:8.
Wilson, H.L. 1910. Gracilariophila, a new parasite on Gracilaria confervoides. Univ.
Calif. Publ. Bot., 4: 75-85.
Wynne, M.J. 1989. The re-instatement of Hydropuntia Montagne (Gracilariaceae, Rhodophyta). Taxon, 38: 476-479.
Xia, B. 1986. On Gracilaria salicornia (C. Agardh) Dawson. Chinese Journal of Oceanology and Limnology, 4:100-107.
Yang, E.C., M.S. Kim, P.J.L. Geraldino, D. Sahoo, J.A. Shin, and S.M. Boo. 2008.
Mitochondrial cox1 and plastid rbcL genes of Gracilaria vermiculophylla
- 53 -
(Gracilariaceae, Rhodophyta). J. Appl. Phycol., 20: 161-168.
Yamamoto, H. 1978. Systematic and anatomical study of the genus Gracilaria in Japan.
Memoirs of the Faculty of Fisheries Hokkaido University, 25: 97-152.
Yamamoto, H. 1986. Congracilaria babae gen. et sp. nov. (Gracilariaceae), an
adelphoparasite growing on Gracilaria salicornia of Japan. Bull. Fac. Fish. Hokkaido.
Univ., 37: 281-290.
- 54 -
국 문 초 록
홍조류 꼬시래기과는 아가로즈를 생산하는 주요 해조류로서 현재
150 여종의 생육이 전 세계적으로 널리 보고되었다. 꼬시래기과는
유용해조류라는 중요성 때문에 생활사, 배양, 분류 등 폭넓은 연구가 진행되어 왔으나 종을 식별하는데 사용하는 뚜렷한 기준형질이 모호하고, 개체간, 지역간 환경에 따른 형태변이가 심하며, 과 내의 많은 종 수로 인하여 분류학적 및 종의 경계에 대한 문제가 많았다. 본 연구는 아시아-태평양지역에 서식하고 있는 유용 홍조류 꼬시래기과의 계통학적 유연관계와 종 다양성을 밝히고, 종의 경계를 명확히 하는 데에 그 목적이 있다. 이를 위해 한국, 일본, 말레이시아 등 7 개국 연안의 조간대 및 조하대에서 채집된
꼬시래기과의 115 개체를 연구재료로 사용하였으며, 색소체 rbcL 과
미토콘드리아 COI 유전자 분석이 이루어졌다. RbcL 1,171 염기쌍과 COI
616 염기쌍이 분석된 결과, 5 종의 잠재종을 포함한 총 22 종의 생육 및
지리적 분포가 확인되었다. RbcL 유전자 분석을 통해 추론된 분자계통
결과에서 Gracilaria, Gracilariopsis, Hydropuntia 속은 단계통을 이루었고,
22 종의 분류학적 위치 및 계통학적 유연관계를 확인하였다.
최대절약(Maximum parsimony), 최대유사 (Maximum likelihood) 방법을
이용한 분석에서는 Hydropuntia 와 G. vermiculophylla 그룹을
포함하는 Gracilaria sensu lato 와 Gracilariopsis 으로 크게 나뉘었으며, 꼬시래기과내의 유전적 변이율은 1.1-14.6%로 나타났다. COI 유전자 분석을
- 55 -
통한 DNA 바코딩 연구결과를 통해 7 종의 Gracilaria, 2 종의 Hydropuntia, 2 종의 Gracilariopsis 를 확인하였다. 0-1.5%의 종내 변이율과 2.8-16.9%
종간 변이율이 관찰되었고, 모든 종이 근린결합분석 (Neighbor joining)을 이용한 계통수에서 각각의 독립적인 계통군을 형성하고 있어 DNA 바코딩 기법이 꼬시래기 종의 식별에 효율적임을 보여주고 있다. 한편, 아시아-태평양지역 7 개국에서의 채집을 통하여 3 종의 미기록종을 확인하였다.
기준생육지인 호주에서만 생육이 보고되어 왔던 G. perplexa 는 일본의
오키나와섬에서 생육을 확인하였고, 태국과 베트남에 보고된 H. fisheri 와
중국의 하이난섬, 필리핀에서 보고된 Gp. heteroclada 는 말레이시아에서의
생육을 확인하였다. 또한 현재까지 보고된 종과는 다소 차이를 나타내는 외부형태와 DNA 염기서열을 가지는 5 개의 잠재종을 확인하였으며, 이에 대한 분류학적 위치 및 계통학적 유연관계를 규명하기 위하여 추가적인 연구가 필요할 것으로 사료된다.
- 56 -
- 57 -