Journal Search Engine
Search Advanced Search Adode Reader(link)
Download PDF Export Citaion korean bibliography PMC previewer
ISSN : 1226-9999(Print)
ISSN : 2287-7851(Online)
Environmental Biology Research Vol.31 No.4 pp.243-274
DOI : https://doi.org/10.11626/KJEB.2013.31.4.243

한국연안해역 와편모조류 시스트 연구에 관한 고찰

윤 양 호*, 신 현 호1
전남대학교 수산해양대학 해양기술학부, 1한국해양과학기술원 남해연구소
한국 연안 해역의 와편모조류 시스트의 연구 현황과 특성을 파악하기 위해 한국 연안과 내만은 물론 제주도 남부와 황해 일부해역을 포함한 동중국해를 대상으로 수행한 45편의 학술연구논문을 분석하였다. 결과 한국의 와편모조류 시스트는 1980년대 후반부터 연구가 시작되었고, 대상 해역은 적조발생 등 부영양화 해역이나 연안개발이 이루어지는 해역에 집중되었으며, 동해를 대상으로 한 연구결과는 없었다. 연구 내용도 단순한 출현 종분석에서 부영양화 진행 등 해양환경의 변화과정 추적, 적조 발생에서 종자군으로서 역할, 세디맨트 트랩을 이용한 시스트의 형성과 변화양상 추적, 수소이온 농도 등 해양환경 변화에 따른 형태변화, 실험실에서 특정 종의 발아 특성 등 국제적 수준의 다양한 연구가 진행되고 있었다. 출현종이나 출현 세포밀도 역시 해역에 따라 매우 다양한 특성을 보인다. 그러나 한국의 와편모조류 시스트 연구는 아직 체계적 연구가 이루어지지 못하고 있고, 단편적이면서 개별적 내용으로 매우 제한적인 연구성격을 나타내었다. 특정 해역에 편중된 연구결과와 특정 연구 연구 집단에 의해 산발적인 연구가 수행되는 경향을 나타내었다. 해양에서 표층퇴적물의 와편모조류 시스트는 표영 환경의 누적지표를 나타내는 것으로 해양환경 변화를 효율적으로 추적하는 데 유용한 지표가 되고 있음을 고려할 때, 국내에서도 조직적인 인력양성과 함께 해양생태계의 구성인자로서 필수 해양생물 분야로 취급하는 연계적인 연구 추진이 절실하게 요구된다.

Summary on the Dinoflagellate Cyst Assemblages of Modern Sediments from Korean Coastal Waters and Adjoining Sea

Yang Ho Yoon*, Hyeon Ho Shin1
Faculty of Marine Technology, College of Fisheries and Ocean Science, Chonnam National University, Yeosu 550-749, Korea
1South Sea Institute, Korea Institute of Ocean Science and Technology, Geoje 656-830, Korea
Received: 2 September 2013, Revised: 22 September 2013, Revision accepted: 23 September 2013

Abstract

To investigate the research state and characteristics of dinoflagellate cyst along theKorean coastal waters, this work analyzed 45 studies conducted in the Korean coasts and basin,the south of Jeju, East China Sea including some part of the Yellow Sea. It was found that theresearch on dinoflagellate cyst began in the later half of the 1980s in Korea, and that the researchareas focused on eutrophied regions where a red tide occurred or on the seas where coastal developmentoccurred. In other words, no research on the East Sea was found. In terms of research contents,there were various studies on analysis of cyst assemblage, tracking of the changes in marine environmentlike process of eutrophication, roles as a seed population in occurrence of a red tide, creationof cyst by use of sediment trip and tracking of its changes, morphological changes by the changeof pH concentration in the hypoxia zone of eutrophied region, germination of a specific species, andother international-level studies. Species composition and cell density also varied in the Koreancoastal waters. However, much research on dinoflagellate cyst in Korea is not systematic, and isfragmentary and individual and hencesuffers from great limitations. The research results so farhave focused only on specific regions, and sporadic research had been conducted by specific researchgroups. Dinoflagellate cyst of surface sediments represents the accumulation index of pelagic environmentin the sea and is a useful index to understand the marine environment efficiently. In conclusion,Korea requires organized human resources and collaborative research on dinoflagellate cystwhich in turn should be considered as a component of marine ecosystem and as an essential aspectof marine biology.

0053-01-0031-0004-2.pdf408.2KB

서 론

 와편모조류는 단세포의 생물로서 해양의 기초생산자에서 규조류 다음으로 중요한 위치를 차지하는 분류군으로 현재 약 2,500종이 알려진다(Taylor et al. 2008; Hoppenrath et al. 2009). 단세포 생물의 일반적인 번식은 핵분열에 의한 무성생식을 기본으로 한다. 와편모조류의 유영세포 역시 무성생식으로 성장하지만, 서식환경이 나빠지면 배우자의 접합으로 유성생식으로 변화한다. 유영세포의 배우자 접합으로 일시적으로는 두 개의 종편모를 가지는 운동성 접합자(planozygote)가 되지만, 얼마 없어 내부에 휴면성 접합자(hypnozygote, resting spore, resting cyst)를 만들어 운동성 접합자의 세포벽 대신에 내구성의 강한 유기물 세포벽을 만들어 운동성 상실은 물론 유영세포와는 형태를 달리하는 휴면포자 (resting cyst))가 되어 해저 표층퇴적층에 침강하여 휴면상태에 들어간다. 이와 같은 휴면상태의 세포를 일반적으로 시스트(cyst)라고 불리고 있다. 휴면포자는 몇 주에서 몇 개월을 해저 표층퇴적층에서 휴면한 다음, 수온 등 외부의 환경 자극에 의해 재차 발아하여 두 개의 종편모를 가지는 운동성 감수모세포(planomeiocyte)로 표영 환경으로 돌아와 무성생식에 의한 유영세포 시기를 보낸다. 다만 아직까지 와편모조류의 많은 종에 대해 휴면포자를 포함한 생활사가 분명하게 밝혀져 있지 않다(Matsuoka 1999). 현재 유영세포로 발견되는 와편모조류의 약 10~20% 정도가 생활사에서 유성생식에 의한 시스트를 만드는 것으로 추정되고 있다(Head 1996; Price and Pospelova 2011; Bonnet et al. 2012).

 와편모조류가 휴면기 세포인 시스트를 만드는 이유는 수권 생태계에서 군집의 생존율을 높이기 위한 전략(Odebrecht 1982; Dale 1983; Walker 1984; Sarjeant et al. 1987; Heiskanen 1993)임과 동시에 유영세포의 군집을 확산시키거나 유전자의 재조합을 위한 생존전략이라 할 수 있다(Anderson 1984; Sarjeant et al. 1987; Matsuoka et al. 2010). 이와 같은 시스트의 형성과 발아에 영향을 주는 인자로는 수온, 질소와 인 등 영양염류, 광량, 염분, 미량금속 원소, 용존산소, 황화수소, 극지방의 얼음 피도와 같은 다양한 무생물 환경과 세균이나 기초생산과 같은 생물학적 환경 등이 알려진다(Bibby and Dodge 1972; Anderson et al. 1987; Ellegaard et al. 1998, Kremp and Anderson 2000; Sgrosso et al. 2001; Yoo and Shin 2004; Matrai et al. 2005; Figueroa et al. 2011; Candel et al. 2012 etc).

 와편모조류 시스트의 생리∙생태학적 특성으로 현생표층퇴적물의 시스트 연구는 약 50년 정도의 짧은 역사임도 불구하고, 와편모조류의 생활사 규명을 위한 생태학적 연구(van Stosch 1973; Walker and Steidinger 1979; Morey-Gains and Ruse 1980; Anderson et al. 1983; Kita et al. 1993; Uchida et al. 1996; Kim and Han 2000; Anderson and Rengefors 2006; Alpermann et al. 2009; Bravo et al. 2012 etc)는 물론, 해수유동, 전선, 용승, 유빙과 수리 등의 물리학적 현상 파악을 위한 생물학적 증거자료와 무산소 수괴나 수질 환경의 지시종 등 해양현상의 이해(Wall et al. 1977; Tyler et al. 1982; Anderson and Keafer 1987; Nehring 1997; Devillers and de Vernal 2000; Gu et al. 2003; Yoo and Shin 2004; Sorrel et al. 2006; Solignac et al. 2008; Elshanawany et al. 2010; de Vernal and Rochon 2011; Bonnet et al. 2012 etc), 그리고 매년 같은 시기에 동일 장소에서 유독/유해성 와편모조류에 의한 적조 발생의 종자군(seed population) 역할과 유독/유해 와편모조류의 분포도 작성, 산업활동의 결과로 발생하는 연안/내만해역의 부영양화 등 환경변화의 추적자로서도 유용한 연구 수단으로 이용된다(Steidinger 1975; Anderson and Keafer 1987; Kim et al. 1990; Dale et al. 1999; Matsuoka 1999, 2001, 2011; Dale 2000, 2001a, b, 2009; Klemp and Anderson 2000; Joyce et al. 2002; Joyce and Pitcher 2004; Wang et al. 2007; Smayda 2008; Shin et al. 2010b; Matsuyama 2012; Usup et al. 2012; Satta et al. 2013 etc). 또한 선박 균형수(ballast water) 등에 의한 유독와편모조류 등 종의 확산과 외래종의 도입(Hallegraeff et al. 1990; Hallegraeff and Bolch 1992; Ichikawa et al. 1992; Hamer et al. 2001; Hong et al. 2003; Patil et al. 2005; Pertola et al. 2006; Bolch and de Salas 2007; Fahnenstiel et al. 2009; Dai et al. 2012 etc)과 고환경을 포함한 기후 변화를 해석하는 수단으로도 유용하게 이용되며(Chalson et al. 1987; Edwards et al. 1991; Wall et al. 1977; Kunz-Pirrung et al. 2001; Pospelova et al. 2006; Grøfjeld et al. 2009; Guiot and de Vernal 2011; Naidu et al. 2012; Price et al. in press etc), 생물진화 및 생존전략, 어패류의 독화에 의한 공중 보건 위생의 위협, 상위 영양단계 생물의 섭식행동, 먹이사슬을 통한 종의 재확산은 물론 생태계 및 해양공간에서의 물질 수송 등의 분야(Wall and Dale 1968; Dale et al. 1978; Anderson 1984; Takeuchi 1985; Hallegraeff et al. 1989; Han et al. 1992; Corrales and Maclean 1995; Nehring, 1996; Anderson 1998; Persson 2000; Harper et al. 2002; Kremp et al. 2003; Lan et al. 2004; Bazzoni et al. 2006; Hoppenrath and Leander 2007; Cox et al. 2008; Persson and Smith 2009a, b; Anderson et al. 2012; Satta et al. 2013 etc) 및 생물지리 분포(Marret and Zonn-eveld 2003; Zonneveld et al. 2013) 등 다양한 분야에서 연구의 대상으로 이용된다.

 국내에서 와편모조류 시스트에 대한 연구는 1980년대초 내만 해역에서 적조발생과 관련하여 시도된 연구보고(Kim et al. 1990; Lee and Yoo 1991; Moon et al. 1993)를 시초로 하여, 최근 다양한 연구가 수행되고 있지만, 매우 편향된 연구에 지금까지의 연구 내용을 정리한 내용이 없다. 따라서 국내의 와편모조류 시스트 연구의 실태와 특성에 대하여 정리하여, 앞으로 와편모조류 시스트 연구를 수행하고자 하는 연구자들에게 도움을 주고자 본고를 정리한다.

재료 및 방법

 한국 연안 해역의 와편모조류 시스트 연구 현황과 특성을 분석하기 위해 제공한 자료는 2013년 8월까지 국내∙외의 전문학술지와 전문도서로 발간된 것만을 대상으로 하였다. 와편모조류 시스트를 대상으로 하여 작성된 학위논문이나 연구보고서가 있지만, 이들은 분석 대상에서 제외하였다. 그리고 대상 해역은 한국의 내만과 연안 해역은 물론 제주도 남부해역을 포함하는 동중국해와 황해을 대상으로 국내 연구자가 참여하여 작성한 학술논문은 대상에 포함시켰다. 분석에 사용한 문헌을 발표 년대 순으로 주된 연구내용과 함께 Table 1에 정리하였다. 국내 전문학회지 25편, 국제전문학회지 16편을 포함하여 45편의 문헌이다.

Table 1. The list of studies on the dinoflagellate cysts in Korean coastal water (KCW)

결과 및 고찰

1. 와편모조류 시스트의 연구변화와 특성

 한국 연안 해역에서 수행된 와편모조류 시스트의 연구는 1982년 한국 동남 연안 해역의 적조모니터링 정점의 표층퇴적물을 대상으로 한 것이 처음이다(Kim et al. 1990). 1982년 2월에서 11월까지 주간 또는 격주로 약 40개의 정점을 대상으로 분석하여 적조원인종이 되는 와편모조류 시스트 6종과 침편모조류 시스트 2종, 모두 8종의 시스트를 동정하고 있다. 이후 1980년대 진해만의 대규모 적조와 관련하여 표층퇴적물에 포함된 시스트연구는 Lee and Yoo (1991)와 Moon et al. (1993) 등의 보고로 1990년대 전반은 진해만과 만산만을 중심으로 하는 극히 제한된 해역에서 출현종에 대한 연구가 수행되었다. 이와 함께 패류독화 원인종으로 알려진 유독 와편모조류인 Alexandrium의 발아와 독조성에 대한 연구가 있다(Kim 1994, 1995). 1990년대 후반에는 대상해역이 진해-마산만에서 통영연안해역과 가막만, 그리고 남해의 개방해역까지 확대되어, 시스트 출현종에 대한 연구가 수행되었다(Lee and Matsuoka 1996; Lee et al. 1998, 1999; Kang et al. 1999). 2000년대에 들어 국내 와편모조류 시스트 연구는 활기를 띠게 되며, 연구 해역도 동해를 제외한 남해와 서해(황해) 일부해역, 그리고 동중국해까지 확대된다. 연구 내용도 출현종 파악에서 한 단계 진행된 발아 환경특성(Kim and Han 2000; Park et al. 2004; Yoo and Shin 2004), 생활사 파악(Kim et al. 2002, 2007; Kim 2009; Park et al. 2009), 주상 퇴적물 표본에 의한 부영양화 변화양상 파악 등 환경 변동 파악(Kim et al. 2003b; Park et al. 2004; Kim et al. 2009) 등으로 연구가 확대되었다. 그리고 최근인 2010년 이후는 양식장 등 특정 해역의 시스트 군집파악과 환경해석 (Pospelova and Kim 2010; Shin et al. 2010), Sediment trap을 이용한 시스트 형성과 변화과정 추적(Shin et al. 2012), 그리고 빈/무산소수역에서의 시스트의 형태 변화(Shin et al. 2013) 등 국제적 수준에 준하는 연구가 수행되고 있다.

 그러나 2000년대 이후 활발한 와편모조류 시스트 연구에도 불구하고 시스트 연구는 적조와 해양개발 등 사회적 이벤트와 관련하여, 해역적으로는 남해안에 집중되어있다(Table 2). 표에서 현재 동해를 대상으로 한 시스트 연구는 발아 실험을 위해 영일만에서 Scrippsiella trochoidea 종을 분리하기 위해 채집한 것(Kim and Han 2000)을 제외하면 전무한 실정이며, 분석 대상 45편의 문헌에서 복수 대상 해역을 중복 포함하더라도 황해를 포함한 서해안 4편, 제주도를 포함하는 동중국해 5편을 제외한 35편은 남해에 집중되어 있다. 특히 진해만, 광양만, 가막만등 적조발생 또는 대규모 임해공업단지 조성에 따른 개발 사업이 이루어지는 남해 중앙부에서 동부해역에 편중된 경향을 나타낸다. 즉 현재 와편모조류 시스트의 연구는 다양한 분야에 응용되고 있고, 새롭게 그 영역을 확대하고 있으나, 국내의 여건은 아직도 제한된 연구 집단에 의해 산발적인 연구가 수행되고 있다고 할 수 있다.

Table 2. Study site and theme of the dinoflagellate cysts in KCW

2. 와편모조류 시스트의 출현종

 현재 한국 연안 해역에 출현이 확인된 시스트 종은 1990년대 초반에는 10종 내외로 매우 제한적이었으나(Kim et al. 1990; Lee and Yoo 1991), 지금까지의 모두를 종합하면 약 90종으로 유영세포의 명칭을 기준으로 하면 61종에 달한다(Table 3). 이와 같이 유영세포와 시스트 기준에서 종 수 차이가 있는 것은 지구에 존재하는 생물군 중 유일하게 와편모조류의 연구에서 유영세포는 생물학자에 의해 시스트는 미고생물 영역인 지질학자에 의해 서로 독립적인 분류기준으로 연구가 수행되었기 때문으로 동일 생물종에 대해 유영세포와 시스트가 각각 다른 종명을 사용하고 있기 때문이다. 이와 같은 현상은 국제식물명명규약에서 용납될 수 없지만, 유영세포의 생활사가 확인된 것은 전체 10% 내외로 시스트와 유영세포 사의의 대응관계가 충분하지 못한 것과 현재 멸종된 2,000여 종의 시스트에 대한 유영세포의 형태 파악은 불가능 하여 화석의 현태로 분류할 수밖에 없다는 두 가지 이유에서 당분간은 동일 종에 대한 두개의 분류체계는 양립될 수밖에 없을 것으로 판단되고 있다(Wall and Dale 1968; Dale 1983; Fensome et al. 1993; Matsuoka and Fukuyo 2000).

Table 3. Species list of dinoflagellate motile cell and cysts from modern sediments in Korean coastal area and adjoining sea

Table 3. Continued

 또한 와편모조류 유영세포는 갑짝스런 담수 유입 등 외부 환경의 일시적인 변화에 대해서도 형태를 변화시켜 일시적으로 해저에 침강하였다가 회복되는 일시성시스트(pellicle cyst, temporary or ecdysal cyst)를 형성하는 것은 잘 알려져 있다(Dale 1977; Walker 1984). 일시성 시스트에서는 운동성 접합자는 관찰되지 않는다. 와편모조류가 유독/유해성 적조원인생물과 표영 환경이 누적지표로 환경 분야 응용뿐만 아니라 독성 단백질 등 일부의 물질은 신약개발이나 기능성 물질 탐색이 중요한 대상이 되고 있어, 와편모조류가 중요 연구 대상 생물군이 되면서 다양한 유영세포에 대해 새로운 시스트의 보고가 매우 활발하다. 이런 과정에 현생 유영세포에 대해 기존의 보고된 많은 시스트에는 시스트가 아닌 일시성시스트임을 지적하는 등(Bravo et al. 2010), 시스트 연구에 안정되지 못한 부분이 많다.

 한국 연안 해역에서 출현이 보고된 90종의 와편모조류 시스트 종에는 특정 연구자에 특정 해역에서만 보고되는 종들이 다수 있어, 발아실험 등 제반 분류학적 검토가 필요한 부분은 있다고 하더라도, 현재 전 세계에서 보고되는 와편모조류 시스트가 150~200종임을 고려하면(Matsuoka and Fukuyo 2000), 국내 출현종은 매우 다양하다고 할 수 있다(Table 3). 표에서 최근 새롭게 발견되는 현생 와편모조류 시스트는 별도의 고생물학적 이름을 붙이지 않고 유영세포의 종명을 시스트 이름을 이용하고 있음을 나타내고 있고, 과거 고생물학적 종명의 시스트를 발아시킬 경우, 현생 유영세포에서 상이한 종이 출현하고 있어, 하나의 유영세포 종명에 다수의 고생물학적 명칭이 나타나고 있는 점에 주의가 필요하다.

 그리고 한국 연안 해역의 시스트 연구가 남해연안에 집중되어 있어, 출현종에 대한 생물ㆍ지리학적 검토가 큰 의미를 나타내는 것은 아니지만, 현재까지의 연구결과를 정리하는 입장에서 출현종별 출현해역과 문헌, 그리고 광학현미경 사진을 첨부하여 정리하여 둔다. 다만 일부 종에 대해서는 현미경 사진을 제시하지 못하였고, 출현종은 시스트를 이용한 부영양화 과정 평가 수단으로 독립영양종과 종속영양종으로 구분하였지만(Dale et al. 1999; Matsuoka 1999, 2001, 2011; Dale 2000, 2001, 2009), 와편모조류 유영세포에서 완전 독립영양을 하는 종은 수종에 지나지 않고 많은 종에서 반독립영양이나 종속영양을 수행하고 있어(Taylor 1987), 단순히 독립영양종과 종속영양종으로 구분하는 것에 대해서도 논쟁이 되고 있다.

AUTOTROPHIC SPECIES

Order GONYAULACALES Taylor 1980
Family CALCOIDINELLACEAE Taylor 1987
(Calciodineloid Group)
Genus Ensiculifera Balech 1967

 Ensiculifera carinata Matsuoka et al. 1990
 Paleontological name: cyst of Ensiculifera carinata Matsuoka et al. 1990 (Plate I-1)
 Distribution: Gamak bay (Shin et al. 2012), western South Sea (Shin et al. 2007)

Plate I. Light microscope photographs of dinoflagellate cysts from Korean coastal waters. 1. cyst of Ensiculifera carinata, 2. cyst of Pentapharsodinium dalei, 3. cyst of Scrippsiella crystallina, 4. cyst of Scrippsiella trochoidea, 5. cyst of Scrippsiella sp., 6. cyst of Alexandrium affinis, 7. cyst of Alexandrium catenatum/tamarense, 8. cyst of Alexandrium minutum, 9. Polysphaeridinium zoharyi, 10. cyst of Alexandrium sp., 11. Spiniferites bentori, 12. Spiniferites elongatus (scale bar=10 μm).

Genus Pentapharsodinium Indelicato et Loeblich III 1986

 Pentapharsodinium dalei (Indelicato et Loeblich III 1986) Montreso et al. 1993
 Paleontological name: Pentapharsodinium dalei (Indelicato et Loeblich III) Montreso et al. 1993 (Plate I-2)
 Synonym: Peridinium faeroense Dale 1977
 Distribution: Tongyeong coasts (Pospelova and Kim 2010)

Genus Scrippsiella Balech ex Loeblich III 1965

 Scrippsiella crystallina Lewis 1991
 Paleontological name: cyst of Scrippsiella crystallina Lewis 1991 (Plate I-3)
 Distribution: Seamangeum coasts (Park et al. 2004; Park and Shim 2011), East China Sea (Cho et al. 2003)

  Scrippsiella trochoidea (Stein) Loeblich III 1976
  Paleontological name: cyst of Scrippsiella trochoidea (Stein) Loeblich III 1976 (Plate I-4)
  Distribution: Yeongil bay (Kim and Han 2000), Jinhae- Masan bay (Lee et al. 1998; Park and Shim 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Kang et al. 1999; Park and Shim 2011), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park et al. 2004; Shin et al. 2010, 2012; Park and Shim 2011; Shin et al. 2013), central South Sea (Lee and Matsuoka 1996; Cho et al. 2003; Park et al. 2005), western South Sea (Shin et al. 2007; Park et al. 2008) Seamangeum coasts (Park et al. 2004; Park and Shim 2011), Yellow Sea (Cho and Matsuoka 2001), East China Sea (Cho and Matsuoka 2001)

 Scrippsiella sp./spp.
 Paleontological name: cyst of Scrippsiella sp. (Plate I-5)
 Distribution: Tongyeong coasts (Kang et al. 1999), Bokdolbada (Shin et al. 2011), central South Sea (Lee and Matsuoka 1996), Yellow Sea (Cho and Matsuoka 2001; Hwang et al. 2009), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001)

Family GONIODOMATACEAE Lindemann 1928
(Gonyaulacoid Group)
Genus Alexandrium Halim 1928

  Alexandrium affinis (Inoue et Fukuyo 1985) Balech 1985
  Paleontological name: cyst of Alexandrium affinis (Inoue et Fukuyo 1985) Balech 1985 (Plate I-6)
  Synonym: Protogonyaulax affinis Inoue et Fukuyo 1985 A. fukuyoi Balech 1985
  Distribution: Jinhae-Masan bay (Kim 1991; Park and Shim 2011), Gamak bay (Lee et al. 1999; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), eastern South Sea (Kim et al. 1990), Bokdolbada (Shin et al. 2011), Seamangeum coasts (Park and Shim 2011), East China Sea (Cho et al. 2001; Kim et al. 2010)

 Alexandrium catenatum (Whedon et Kofoid) Balech 1985/A. tamarense (Lebour) Balech 1992
 Paleontological name: cyst of Alexandrium catenatum/tamarense complex (Plate I-7)
 Synonym: Gonyaulax catenatum Whedon et Kofoid 1936
                  Gessnerium catenatum (Whedon et Kofoid) Loeblich III et L Loeblich 1979
                  Protogonyaulax catenatum (Whedon et Kofoid) Taylor 1979
                 Gonyaulax tamarensis Lebour 1925
                 G. tamarensis var. excavata Braarud 1945
                 G. excavata (Braarud) Balech 1971
                 Gessnerium tamarensis (Lebour) Loeblich III et L. Loeblich 1979
                 Protogonyaulax tamarensis (Lebour) Taylor 1979
                A. excavata (Braarud) Balech et Tangen 1985

 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Lee and Yoo 1991; Moon et al. 1993; Lee et al. 1998; Park and Shim 2011), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Kang et al. 1999; Pospelova and Kim 2010; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), Yellow Sea (Cho and Matsuoka 2001), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2004)

 Alexandrium minutum Halim 1960
 Paleontological name: cyst of Alexandrium minutum Halim 1960 (Plate I-8)
 Synonym: A. ibericum Balech 1985
 Distribution: Gamak bay (Park and Shim 2011), Seamangeum coasts (Park and Shim 2011), East China Sea (Kim et al. 2010)

 Alexandrium sp.
 Paleontological name: cyst of Alexandrium sp. (Plate I-10).
 Distribution: Jinhae-Masan bay (Lee et al. 1998), Gwangyang bay (Kim et al. 2003a, b), central South Sea (Lee and Matsuoka 1996; Cho et al. 2003), Seamangeum coasts (Park et al. 2004), Yellow Sea (Cho and Matsuoka 2001; Hwang et al. 2009), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003)
 Remarks: 문헌에 따라서는 Alexandrium속의 시스트를 종까지 분류하지 않고 Alexandrium sp./spp.로 포괄적 표현을 하거나, 타원형, 구형 등으로 전체적인 형태만을 표현하고 있음.

Genus Pyrodinium Plate 1906

 Pyrodinium bahamense var. compressum (Böhm) Steidinger et al. 1980
 Paleontological name: Polysphaeridinium zoharyi (Rossignol) Bujak et al. 1980
 Distribution: Tongyeong coasts (Pospelova and Kim 2010), Yellow Sea (Hwang et al. 2009)

Family GONYAULACEAE Lindemann 1928
(Gonyaulacoid Group)
Genus Gonyaulax Diesing 1866

 Gonyaulax digitalis (Pouchet) Kofoid 1991
 Paleontological name: Spiniferites bentori (Rossigno) Wall et Dale 1970 (Plate I-11)
 Synonym: G. digitale Lemmerman 1899
                   Protoperidinium digitale Pouchet 1883

 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Moon et al. 1993; Lee et al. 1998; Park and Shim 2011), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Park and Shim 2011; Shin et al. 2011), Tongyeong coasts (Pospelova and Kim 2010), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Yellow Sea (Cho and Matsuoka 200), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003)

 Gonyaulax elongata (Reid) Ellegaard et al. 2003
 Paleontological name: Spiniferites elongatus Reid 1974 (Plate I-12)
 Distribution: Jinhae-Masan bay (Kim 1991; Moon et al. 1993; Lee and Matsuoka 1996; Lee et al. 1998; Park and Shim 2011), Gwangyang bay (Kim et al. 2003a), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011), Yeoja bay (Shin et al. 2010a), Tongyeong coasts (Kang et al. 1999; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), Yellow Sea (Cho and Matsuoka 2001), East China Sea (Cho and Matsuoka 2001)
 Remarks: 많은 문헌에서 유영세포의 명칭을 Gonyaulax spinifera complex로 표현하고 있음.

 Gonyaulax membranaceus (Rossignol) Ellegaard et al. 2003
 Paleontological name: Spiniferites membranaceus (Rossignol) Sarjeant 1970 (Plate II-1)
 Distribution: Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Park and Yoon 2003; Park et al. 2004), Tongyeong coasts ( (Pospelova and Kim 2010), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001)
 Remarks: 많은 문헌에서 유영세포의 명칭을 Gonyaulax spinifera complex로 표현되고 있음.

Plate II. Light microscope photographs of dinoflagellate cysts from Korean coastal waters. 1. Spiniferites membranaceus, 2. Spiniferites bulloideus, 3. Ataxiodinium choane, 4. Nematosphaeropsis labyrinthus, 5. Nematosphaeropsis sp., 6. Spiniferites hypercanthus, 7. Spiniferites mirabilis, 8. Spiniferites ramosus, 9. Bitectatodinium spongium, 10. Impagidinium aculeatum, 11. Spiniferites delicatus, 12. Lingulodinium machaerophorum.

 Gonyaulax verior Sournia 1973
 Paleontological name: cyst of Gonyaulax verior Sournia 1973
 Synonym: Amylax diacantha Meunier 1919
                  Gonyaulax diacantha sensu Schiller 1937
 Distribution: Gamak bay (Lee et al. 1999; Park and Shim 2011), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), Yellow Sea (Hwang et al. 2009)

 Gonyaulax scrippsae Kofoid 1911
 Paleontological name: Spiniferites bulloideus (Deflandre et Cookson) Sarjeant 1905 (Plate II -2)
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Lee and Matsuoka 1996; Lee et al. 1998; Park and Shim 2011; Shin et al. 2011), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a), Geoje coast (Shin et al. 2007), Tongyeong coasts (Kang et al. 1999; Pospelova and Kim 2010; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park et al. 2004), Yellow Sea (Cho and Matsuoka 2001; Hwang et al. 2009), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003; Cho et al. 2004; Kim et al. 2010)

 Gonyaulax spinifera (Claparede et Lachmann) Diesing 1866 complex
 Paleontological name: Ataxiodinium choane Reid 1974 (Plate II-3)
 Synonym: Peridinium spinifera Claparede et Lachmann 1859
 Distribution: Tongyeong coasts (Pospelova and Kim 2010)

 Gonyaulax spinifera complex
 Paleontological name: Nematosphaeropsis labyrinthus (Ostenfeld 1903) Reid 1974 (Plate II-4)
 Distribution: East China Sea (Kim et al. 2010)

 Gonyaulax spinifera complex
 Paleontological name: Nematosphaeropsis sp. (Plate II-5)
 Distribution: Tongyeong coasts (Pospelova and Kim 2010)

 Gonyaulax spinifera complex
 Paleontological name: Spiniferites hypercanthus (Deflandre et Cookson) Cookson et Eisenack 1974 (Plate II-6)
 Distribution: Busan harbor (Kim et al. 2005), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Park and Yoon 2003; Park et al. 2004; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park and Shim 2011), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003; Cho et al. 2004; Kim et al. 2010)

 Gonyaulax spinifera complex
 Paleontological name: Spiniferites mirabilis (Rossignol) Sarjeant 1970 (Plate II-7)
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Moon et al. 1993), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2012), Geoje coast (Shin et al. 2007), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park and Shim, 2011), Yellow Sea (Cho and Matsuoka, 2001), East China Sea (Cho and Matsuoka, 2001; Cho et al. 2001; Cho et al. 2003; Cho et al. 2004; Kim et al. 2010)

 Gonyaulax spinifera complex
 Paleontological name: Spiniferites ramosus (Ehrenberg) Mantell 1854 (Plate II-8)
 Distribution: Busan harbor (Kim et al. 2005), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Park and Yoon 2003; Park et al. 2004; Park and Shim 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Pospelova and Kim 2010; Park and Shim 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park and Shim 2011), Yellow Sea (Cho and Matsuoka 2001), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003; Cho et al. 2004; Kim et al. 2010)

 Gonyaulax sp.
 Paleontological name: Bitectatodinium spongium (Zonneveld) Zonneveld et Jurkschat 1999 (Plate II-9)
 Distribution: Tongyeong coasts ( (Pospelova and Kim 2010) Gonyaulax sp.
 Paleontological name: Impagidinium aculeatum (Well) Lentin et Williams 1981 (Plate II-10)
 Distribution: Tongyeong coasts ( (Pospelova and Kim 2010)

 Gonyaulax sp.
 Paleontological name: Impagidinium sp.
 Distribution: Tongyeong coasts ( (Pospelova and Kim 2010), East China Sea (Cho et al. 2003; Kim et al. 2010)

 Gonyaulax sp.
 Paleontological name: Spiniferites delicatus Reid 1974 (Plate II-11)
 Distribution: Gwangyang bay (Kim et al. 2003a, b), Jinhae- Masan bay (Moon et al. 1993), Gamak bay (Park et al. 2004; Shin et al. 2010b), Yeoja bay (Shin et al. 2010a), Tongyeong coasts (Kang et al. 1999), central South Sea (Cho et al. 2003; Park et al. 2005), Yellow Sea (Cho and Matsuoka 2001), East China Sea (Cho and Matsuoka 2001; Cho et al. 2004; Kim et al. 2010)

 Gonyaulax sp.
 Paleontological name: Spiniferites spp.
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan Bay (Moon et al. 1993), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Kang et al. 1999; Pospelova and Kim 2010), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Lee and Matsuoka 1996; Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), Yellow Sea (Cho and Matsuoka 2001; Hwang et al. 2009), East China Sea (Cho and Matsuoka 2001; Cho et al. 2003; Cho et al. 2004; Kim et al. 2010)

Genus Lingulodinium (Stein) Dodge 1989

 Lingulodinium polyedrum (Stein) Dodge 1989
 Paleontological name: Lingulodinium machaerophorum (Deflandre et Cookson) Wall 1967 (Plate II-12)
 Synonym: Gonyaulax polyedra Stein 1833
 Distribution: Jinhae-Masan bay (Kim 1991; Moon et al. 1993; Park and Shim 2011), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Pospelova and Kim 2010), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), Yellow Sea (Hwang et al. 2009), East China Sea (Cho et al. 2001; Cho et al. 2003; Cho et al. 2004; Kim et al. 2010)

Genus Protoceratium Bergh 1881

 Protoceratium reticulatum (Claparde et Lachmann) Bütschlii 1885
 Paleontological name: Operculodinium centrocarpum sensu Wall et Dale 1966 (Plate III-1)
 Synonym: Gonyaulax grindleyi Reineeke 1967
 Distribution: Jinhae-Masan bay (Kim 1991; Moon et al. 1993; Lee et al. 1998; Park and Shim 2011), Gwangyang bay (Kim et al. 2003a), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Pospelova and Kim 2010), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), Yellow Sea (Cho and Matsuoka 2001; Hwang et al. 2009), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003; Cho et al. 2004; Kim et al. 2010)

Plate III. Light microscope photographs of dinoflagellate cysts from Korean coastal waters. 1. Operculodinium centrocarpum sensu, 2. Operculodinium israelianum, 3. cyst of Cochlodinium sp., 4. cyst of Gymnodinium catenatum, 5. cyst of Pheopolykrikos hartmannii, 6. Tuberculodinium vancampoae.

 Protoceratium sp. cf. reticulatum (Claparde et Lachmann) Bütschli 1885
 Paleontological name: Operculodinium israelianum (Rossignol) Wall 1967 (Plate III-2)
Distribution: Jinhae-Masan bay (Moon et al. 1993), Gamak bay (Park and Yoon 2003; Park et al. 2004), Geoje coast (Shin et al. 2007), Tongyeong coasts (Pospelova and Kim 2010), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003; Kim et al. 2010)
 Remarks: 유영세포에서는 종 수준까지 동정이 되고 있지 못함.

 Protoceratium sp.
 Paleontological name: Operculodinium crassum Harland 1979
 Distribution: East China Sea (Cho et al. 2003)
 Remarks: 유영세포에서는 종 수준까지 동정이 되고 있지 못함.

Order GYMNODINIALES Lemmermann 1910
Family GYMNODINIACEAE Lankester 1885
(Gymnodiniod Group)
Genus Cochlodinium Schütt 1896

 Cochlodinium polykrikoides. Margalef 1961
 Paleontological name: cyst of Cochlodinium polykrikoides Margalef 1961
 Synonym: C. heterolobatum Silva 1967
 Distribution: Jinhae-Masan bay (Park and Shim 2011), Tongyeong coasts (Pospelova and Kim 2010)
 Remarks: 휴면포자인 cyst 존재에 대해 오랜 기간 토론 대상이 되었던 종으로 아직도 명확하지 못함 점은 남아있지만, 최근 새로운 연구결과(Matsuoka et al. 2010; Tang et al. 2012)를 참고로 하여 본 종의 시스트명을 사용함.

 Cochlodinium sp.
 Paleontological name: cyst of Cochlodinium sp. (Plate III-3)
 Distribution: Jinhae-Masan bay (Lee et al. 1998), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Park and Shim 2011; Shin et al. 2012), Tongyeong coasts (Park and Shim 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Park et al. 2005), eastern South Sea (Kim et al. 1990), Seamangeum coasts (Park and Shim 2011), East China Sea (Cho and Matsuoka 2001)
 Remarks: cyst of Cochlodinium polykrikoides 존재가 분명하여 짐에 따라 C. polykrikoides 시스트로 소속을 변경하어야 할 내용도 있지만, cyst of Cochlodinium sp.가 별도 종으로 존재하고 있어, 원 논문의 표현을 그대로 사용하고 있음.

Genus Gymnodinium Stein 1878

 Gymnodinium catenatum Graham 1943
 Paleontological name: cyst of Gymnodinium catenatum Graham 1943 (Plate III-4)
 Distribution: Gwangyang bay (Kim et al. 2003a), Gamak bay (Park and Yoon 2003; Park et al. 2004; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Pospelova and Kim 2010; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), Yellow Sea (Hwang et al. 2009),

 Gymmodinium impudicum (Fraga et Bravo) Hansen et Moestrup 2000
 Paleontological name: cyst of Gyrodinium impudicum (Fraga et Bravo) Hansen et Moestrup
 Synonym: Gyrodinium impudicum Fraga et Bravo 1995
 Distribution: Tongyeong coasts (Park and Shim 2011), Seamangeum coasts (Park and Shim 2011)

 Gymnodinium sp.
 Paleontological name: cyst of Gymnodinium sp.
 Distribution: Jinhae-Masan bay (Lee and Matsuoka 1996; Lee et al. 1998), Tongyeong coasts (Kang et al. 1999), Gamak bay (Lee et al. 1999)

Genus Gyrodinium Kofoid et Swezy 1921

 Gyrodinium instriatum Freudenthal et Lee 1963
 Paleontological name: cyst of Gyrodinium instriatum Freudenthal et Lee 1963
 Distribution: Jinhae-Masan bay (Kim 1991; Moon et al. 1993), Gamak bay (Park and Yoon 2003; Park et al. 2004), Eastern South Sea (Kim et al. 1990), Seamangeum coasts (Park and Shim 2011)

 Gyrodinium sp.
 Paleontological name: cyst of Gyrodinium sp.
 Distribution: Jinhae-Masan bay (Lee and Yoo 1991; Lee et al. 1998), Tongyeong coasts (Kang et al. 1999), Gamak bay (Lee et al. 1999)

Genus Pheopolykrikos Chatton 1933

 Pheopolykrikos hartmannii (Zimmermann) Matsuoka et Fukuyo 1986
 Paleontological name: cyst of Pheopolykrikos hartmannii (Zimmermann) Matsuoka et Fukuyo 1986 (Plate III-5)
 Synonym: Polykrikos hartmannii Zimmermann 1930
 Distribution: Jinhae-Masan bay (Kim 1991; Moon et al. 1993; Lee et al. 1998; Park and Shim 2011), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Kang et al. 1999; Pospelova and Kim 2010; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), eastern South Sea (Kim et al. 1990), Seamangeum coasts (Park et al. 2004; Park and Shim 2011)

Family PYROCYSTACEAE (Schütt) Lemmerman 1899 (Tuberculodinoid Group)
Genus Pyrophacus Murray ex Lemmerman 1899

 Pyrophacus horologium Stein 1883
 Paleontological name: cyst of Pyrophacus horologium (?)
 Distribution: Jinhae-Masan bay (Kim 1991)
 Remarks: 시스트 형성 여부가 확실하지 않음.

 Pyrophacus steinii (Schiller) Wall et Dale 1971
 Paleontological name: Tuberculodinium vancampoae (Rossignol) Wall 1967 (Plate III-6)
 Synonym: Pyrophacus horologium var. steinii Schille 1935
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Lee and Yoo 1991; Moon et al. 1993; Lee et al. 1998), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Pospelova and Kim 2010), western South Sea (Shin et al. 2007), central South Sea (Lee and Matsuoka 1996; Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), Yellow Sea (Cho and Matsuoka 2001; Hwang et al. 2009), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003; Cho et al. 2004; Kim et al. 2010)

 Pyrophacus sp.
 Paleontological name: Tuberculodinium sp.
 Distribution: East China Sea (Cho and Matsuoka 2001)

Heterotrophic Species
Order PERIDINIALES Haeckel 1984
Family KOLKWITZIELLACEAE Lindemann 1928 (Diplosaliod Group)
Genus Diplopelta Stein ex Jørgensen 1912

 Diplopelta parva (Abe) Matsuoka 1988
 Paleontological name: cyst of Diplopelta parva (Abe) Matsuoka 1988
 Synonym: Dissodinium parvum Abe 1941
 Distribution: Gamak bay (Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011), Geoje coast (Shin et al. 2007), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), Yellow Sea (Cho and Matsuoka 2001), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003)

Genus Diplosalis Bergh 1881

 Diplopsalis lenticula Bergh 1881
 Paleontological name: cyst of Diplosalis lenticula Bergh 1881
 Synonym: Peridiniopsis lentacula (Bergh) Starmach 1974
                   Diplosalis lenticulata Stein 1883
                   Glenodinium lentacula (Bergh) Schiller
                   Dissodinium lentacula (Bergh) Loebliech III 1973
Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Lee et al. 1998; Park and Shim 2011), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Park and Shim 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Kang et al. 1999; Park and Shim 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003), Seamangeum coasts (Park and Shim 2011), Yellow Sea (Hwang et al. 2009), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003; Cho et al. 2004)

 Diplopsalis sp.
 Paleontological name: Dulbridinium sp.
 Distribution: Tongyeong coasts (Pospelova and Kim 2010)

Genus Gotoius Abe ex Matsuoka 1988

 Gotoius abei Matsuoka 1988
 Paleontological name: cyst of Gotoius abei Matsuoka 1988
 Distribution: Seamangeum coasts (Park and Shim 2011)

Genus Oblea Balech ex Loeblich Jr. et Loeblich III 1966

 Oblea acanthocysta Kawami et al. 2006
 Paleontological name: cyst of Oblea acanthocysta Kawami et al. 2006 (Plate IV-1)
 Distribution: Gamak bay (Shin et al. 2012), Yeoja bay (Shin et al. 2010a)

Plate IV. Light microscope photographs of dinoflagellate cysts from Korean coastal waters. 1. cyst of Oblea acanthocysta, 2. Dulbridinium caperatum, 3. cyst of Protoperidinium americanum, 4. Brigantedinium cariacoense, 5. Votadinium spinosum, 6. Stelladinium stellatum, 7. Brigantedinium simplex, 8. Selenopempbix quanta, 9. Brigantedinium irregulare, 10. Xandarodinium xanthum, 11. cyst of Protoperidinium latissinum, 12. Lejeunecysta sabrina.

Genus Preperidium Mangin 1913

 Preperidinium meunieri (Pavillard), Elbrächter 1993
 Paleontological name: Dulbridinium caperatum Reid 1977 (Plate IV-2)
 Synonym: Peridinium paulsenii Mangin 1911
                   Diplopsalis minor (Paulsen) Lindermann 1927
                   Zygabikodinium lenticulatum Loeblich III Jr et Loeblich III 1970
                   Peridinium lenticulatum Mangin 1911
                   P. paulsenii Mangin 1911 
                  Diplopsalis lenticula f. minor Paulsen
Distribution: Jinhae-Masan bay (Kim 1991; Lee and Yoo 1991; Moon et al. 1993; Lee et al. 1998; Park and Shim 2011), Gwangyang bay (Kim et al. 2003a), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Geoje coast (Shin et al. 2007), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Tongyeong coasts (Kang et al. 1999; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), central South Sea (Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), East China Sea (Cho and Matsuoka 2001)

Family PERIDINIACEAE Ehrenberg 1831 (Diplosaliod Group)
Genus Heterocapsa Stein 1883

 Heterocapsa triquetra (Ehrenberg) Stein 1883
 Paleontological name: cyst of Heterocapsa triquetra (Ehrenberg) Stein 1883
 Distribution: Jinhae-Masan bay (Kim 1991), estern South Sea (Kim et al. 1990),
 Remarks: 시스트 형성 여부가 확실하지 않음.

Genus Peridinium Ehrenberg 1832

 Peridinium limbatum (Stokes) Lemmerman 1990
 Paleontological name: cyst of Peridinium limbatum (Stokes) Lemmerman 1990
 Synonym: Protoperidinium limbatum Stockes 1888
 Distribution: Tongyeong coasts (Pospelova and Kim 2010)

Genus Protoperidinium Taylor 1987

 Protoperidinium americanum (Gran et Braarud) Balech 1974
 Paleontological name: cyst of Protoperidinium americanum (Gran et Braarud) Balech 1974 (Plate IV-3)
 Synonym: Peridinium americanum Gran et Braarud 1935
 Distribution: Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Kang et al. 1999), Bokdol-bada (Shin et al. 2011), central South Sea (Cho et al. 2003; Park et al. 2005), western South Sea (Shin et al. 2007), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), East China Sea (Cho and Matsuoka 2001; Cho et al. 2004)

 Protoperidinium avellana (Meunier) Balech 1974
 Paleontological name: Brigantedinium cariacoense (Wall) Lentin et Williams 1993 (Plate IV-4)
 Synonym: Peridinium avellana (Meunier) Lebour 1925
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Park and Shim 2011), Gwangyang bay (Kim et al. 2003a), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2011), Tongyeong coasts (Pospelova and Kim 2010), Bokdol-bada (Shin et al. 2011), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park and Shim 2011), Yellow Sea (Cho and Matsuoka 2001), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2004)

 Protoperidinium claudicans (Paulsen) Balech 1974
 Paleontological name: Votadinium spinosum Reid 1977 (Plate IV-5)
 Synonym: Peridinium claudicans Paulsen 1907
 Distribution: Jinhae-Masan bay (Moon et al. 1993), Gwangyang bay (Kim et al. 2003a), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Tongyeong coasts (Kang et al. 1999; Pospelova and Kim 2010; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), East China Sea (Cho and Matsuoka 2001)

 Protoperidinium compressum (Abe) Balech 1974
 Paleontological name: Stelladinium abeii Matsuoka 1985
                                    Stelladinium reidii Bradford 1975
                                    Stelladinium stellatum (Wall et dale 1968) Reid 1977 (Plate IV-6)
 Synonym: Peridinium compressum (Abe) Nie
                   Pongruentinium compressum Abe 1927
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Moon et al. 1993; Lee and Matsuoka 1996; Lee et al. 1998; Park and Shim 2011), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Pospelova and Kim 2010), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Park et al. 2005), Seamangeum coasts (Park and Shim 2011), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003; Cho et al. 2004)

 Protoperidinium conicoides (Paulsen) Balech 1973
 Paleontological name: Brigantedinium simplex Wall 1965 ex Lentin et Williams 1993 (Plate IV-7)
 Synonym: Peridinium conicoides Paulsen 1907
 Distribution: Busan harbor (Kim et al. 2005), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a), Tongyeong coasts (Pospelova and Kim 2010), Geoje coast (Shin et al. 2007), Bokdol-bada (Shin et al. 2011), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park and Shim 2011), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2004)

 Protoperidinium conicum (Gran) Balech 1974
 Paleontological name: Selenopempbix quanta (Bradford 1975) Matsuoka 1985 (Plate IV-8)
 Synonym: Peridinium conicum (Gran) Ostenfeld et Schmidt 1905
                   Peridinium divergens var. conica Gran 1900
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Lee and Yoo 1991; Moon et al. 1993; Lee et al. 1998; Park and Shim 2011), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Tongyeong coasts (Kang et al. 1999; Pospelova and Kim 2010; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Lee and Matsuoka 1996; Park et al. 2005), Seamangeum coasts (Park et al. 2004), Yellow Sea (Hwang et al. 2009), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003; Cho et al. 2004; Kim et al. 2010)

 Protoperidinium denticulatum (Gran et Braarud) Balech 1974
 Paleontological name: Brigantedinium irregulare Matsuoka 1987 (Plate IV-9)
 Synonym: Protoperidinium clavus Abe 1936
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Moon et al. 1993; Park and Shim 2011), Gamak bay (Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Bokdol-bada (Shin et al. 2011), central South Sea (Park et al. 2005), Seamangeum coasts (Park and Shim 2011), East China Sea (Cho et al. 2004)

 Protoperidinium divaricatum (Meunier) Parke et Dodge 1976
 Paleontological name: Xandarodinium xanthum Reid 1977 (Plate IV-10)
 Synonym: P. gainii (Dangeard) Balech 1974
                  Protoperidinium divaricatum (Meunier) Balech 1988
 Distribution: Gamak bay (Park and Yoon 2003; Park et al. 2004), Tongyeong coasts (Pospelova and Kim 2010), Bokdol- bada (Shin et al. 2011), western South Sea (Shin et al. 2007), Seamangeum coasts (Park and Shim 2011), East China Sea (Kim et al. 2010)

 Protoperidinium latissimum (Kofoid) Balech 1974
 Paleontological name: cyst of Protoperidinium latissinum (Kofoid) Balech 1974 (Plate IV-11)
 Synonym: Protoperidinium exiquipes (Mangin) Dodge
                   Protoperidinium pentagonoides Balech 1949
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Lee and Yoo 1991; Lee et al. 1998), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Kang et al. 1999; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park and Shim 2011), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2004)

 Protoperidinium leonis (Pavillard) Balech 1974
 Paleontological name 1: Lejeunecysta sabrina (Reid 1977) Bujak 1984 (Plate IV-12)
 Synonym: Peridinium leonis Pavillard 1916
 Distribution: Jinhae-Masan bay (Lee et al. 1998), Tongyeong coasts (Pospelova and Kim 2010), central South Sea (Cho et al. 2003), East China Sea (Cho and Matsuoka 2001)
 Paleontological name 2: Quinquecuspis concreta Reid 1977(Plate V-1)
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Lee and Yoo 1991; Moon et al. 1993; Park and Shim 2011), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Pospelova and Kim 2010; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park et al. 2004), East China Sea (Cho and Matsuoka 2001; Cho et al. 2003; Kim et al. 2010)

Plate V. Light microscope photographs of dinoflagellate cysts from Korean coastal waters. 1. Quinquecuspis concreta, 2. Votadinium calvum, 3. cyst of Protoperidinium minutum, 4. Brigantedinium majusculum, 5~6. Trinovantedinium applanatum, 7. Selenopemphix alticinctum, 8. cyst of Protoperidinium thorianum, 9. cyst of Protoperidinium sp., 10. rigantedinium auranteum, 11. Brigantedinium asymmetricum, 12. Brigantedinium grande.

 Protoperidinium minutum (Kofoid) Loeblich III 1970
 Paleontological name: cyst of Protoperidinium minutum (Kofoid) Loeblich III 1970 (Plate V-2)
 Synonym: Peridinium minutum Kofoid 1907
 Distribution: Jinhae-Masan bay (Kim 1991; Lee et al. 1998; Park and Shim 2011), Tongyeong coasts (Kang et al. 1999; Park and Shim 2011), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Bokdol-bada (Shin et al. 2011)

 Protoperidinium oblongum (Aurivillius) Parke et Dodge 1976
 Paleontological name: Votadinium calvum Reid 1977 (Plate V-3)
 Synonym: Peridinium oblongum Aurivillius 1898
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Lee and Yoo 1991; Moon et al. 1993; Lee et al. 1998; Park and Shim 2011), Tongyeong coasts(Kang et al. 1999; Pospelova and Kim 2010; Park and Shim 2011), Geoje coast (Shin et al. 2007), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Lee and Matsuoka 1996; Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), Yellow Sea (Hwang et al. 2009), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003; Kim et al. 2010)

 Protoperidinium petagonum (Gran) Balech 1974
 Paleontological name 1: Brigantedinium majusculum Reid 1977 ex Lentin et Williams 1993 (Plate V-4)
 Synonym: Peridinium petagonum Gran 1902
                   Protoperidinium expensum Abe 1981
                   P. parapetagonum Wang 1936
                   P. sinuosum Lemmerman 1905
 Distribution: Gamak bay (Shin et al. 2012), central South Sea (Cho et al. 2003), Yellow Sea (Cho and Matsuoka 2001), East China Sea (Cho and Matsuoka 2001; Cho et al. 2003)
 Paleontological name 2: Trinovantedinium applanatum (Bradford 1977) Bujak et Davies 1983 (Plate V-5, 6)
 Distribution: Gamak bay (Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Bokdolbada (Shin et al. 2011)
 Paleontological name 3: Trinovantedinium capitatum Reid 1977
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Lee and Yoo 1991; Moon et al. 1993; Lee et al. 1998; Park and Shim 2011), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Shin et al. 2010b; Park and Shim 2011), Geoje coast (Shin et al. 2007), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2004; Kim et al. 2010)
 Remarks: 유영세포의 한 종명에 다소 다른 형태를 나타내는 휴면세포, 시스트가 보고되고 있어, Brigantedinium majusculum (Baula et al. 2011), Trinovantedinium applanatum (Marrel and Zonneveld, 2003), Trinovantedinium capitatum (Baula et al. 2011) 등 다양한 고생물명(화석명)으로 표현되고 있음.

 Protoperidinium stellatum (Wall) Balech 1994
 Paleontological name: cyst of Protoperidinium stellatum (Wall in Wall et dale 1968) Rocho et al. 1999
 Synonym: Peridinium stellatum Wall 1968
 Distribution: Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Shin et al. 2010b; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Tongyeong coasts (Pospelova and Kim 2010), Bokdol-bada (Shin et al. 2011), East China Sea (Cho and Matsuoka 2001)

 Protoperidinium subinerme (Paulsen) Loeblich III 1970
 Paleontological name: Selenopemphix nephroides (Benedek 1972) Benedek et Sarjeant 1981 Selenopemphix alticinctum (Bradford)
Matsuoka in Nehring 1997 (Plate V-7)
 Synonym: Peridinium subinerme Paulsen 1904
 Distribution: Jinhae-Masan bay (Moon et al. 1993; Lee and Matsuoka 1996; Lee et al. 1998), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Kang et al. 1999; Pospelova and Kim 2010; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park and Shim 2011), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2004; Kim et al. 2010)

 Protoperidinium thorianum (Paulsen) Balech 1974
 Paleontological name: cyst of Protoperidinium thorianum (Paulsen) Balech 1974 (Plate V-8)
 Synonym: Peridinium thorianum Paulsen 1905
 Distribution: Jinhae-Masan bay (Park and Shim 2011), Seamangeum coasts (Park and Shim 2011)

 Protoperidinium spp.
 Paleontological name: cyst of Protoperidinium sp./spp. (Plate V-9)
 Distribution: Jinhae-Masan bay (Lee et al. 1998), Tongyeong coasts (Kang et al. 1999; Pospelova and Kim 2010), Gamak bay (Lee et al. 1999), central South Sea (Lee and Matsuoka 1996; Cho et al. 2003), Seamangeum coasts (Park et al. 2004), Yellow Sea (Hwang et al. 2009), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003)

 Protoperidinium sp.
 Paleontological name: Brigantedinium auranteum Reid 1977 (Plate V-10)
 Distribution: East China Sea (Cho and Matsuoka 2001; Cho et al. 2001)

 Protoperidinium sp.
 Paleontological name: Brigantedinium asymmetricum Matsuoka 1987 (Plate V-11)
 Distribution: Gamak bay (Shin et al. 2011), Yeoja bay (Shin et al. 2010a), Bokdol-bada (Shin et al. 2011)

 Protoperidinium sp.
 Paleontological name: Brigantedinium grande Matsuoka 1987 (Plate V-12)
 Distribution: Gamak bay (Shin et al. 2012)

 Protoperidinium spp.
 Paleontological name: Brigantedinium sp. (Plate VI-1)
 Distribution: Busan harbor (Kim et al. 2005), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Park and Yoon 2003; Park et al. 2004; Shin et al. 2010b; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Pospelova and Kim 2010; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park and Shim 2011), Yellow Sea (Cho and Matsuoka 2001), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2003; Cho et al. 2004; Kim et al. 2010)

Plate VI. Light microscope photographs of dinoflagellate cysts from Korean coastal waters. 1. Brigantedinium sp. 2. Echinidinium aculenatum, 3. Echinidinium delicatum, 4. Echinidinium sp. cf. delicatum, 5. Echinidinium granulatum, 6. Echinidinium sp., 7. Islandinium brevispinosum, 8. Islandinium minutum, 9. Stelladinium robustum, 10. Trinovantedinium pallidifulvum, 11~12. cyst of Polykrikos kofoidii, 13. cyst of Polykrikos schwartzii.

 Protoperidinium sp.
 Paleontological name: Echinidinium aculenatum Zonneveld 1997 (Plate VI-2)
 Distribution: Gamak bay (Shin et al. 2010b), Yeoja bay (Shin et al. 2010a), Tongyeong coasts (Pospelova and Kim 2010)

 Protoperidinium sp.
 Paleontological name: Echinidinium delicatum Zonneveld 1997 (Plate VI-3)
 Distribution: Tongyeong coasts (Pospelova and Kim 2010)

 Protoperidinium sp.
 Paleontological name: Echinidinium sp. cf. delicatum Zonneveld 1997 (Plate VI-4)
 Distribution: Tongyeong coasts (Pospelova and Kim 2010)

 Protoperidinium sp.
 Paleontological name: Echinidinium granulatum Zonneveld 1997 (Plate VI-5)
 Distribution: Tongyeong coasts (Pospelova and Kim 2010)

 Protoperidinium spp.
 Paleontological name: Echinidinium sp./spp. (Plate VI-6)
 Distribution: Tongyeong coasts (Pospelova and Kim 2010)

 Protoperidinium sp.
 Paleontological name: Islandinium brevispinosum Pospelova et head 2002 (Plate VI-7)
 Distribution: Tongyeong coasts (Pospelova and Kim 2010)

 Protoperidinium sp.
 Paleontological name: Islandinium minutum (Harland et Reid in Harland et al. 1980) Head et al. 2001 (Plate VI-8)
 Distribution: Tongyeong coasts (Pospelova and Kim 2010)

 Protoperidinium sp.
 Paleontological name: Lejeunecysta aliva (Reid 1977) Turon et Londeix 1988
 Distribution: Tongyeong coasts (Pospelova and Kim 2010)

 Protoperidinium sp.
 Paleontological name: Selenopemphix sp.
 Distribution: East China Sea (Cho and Matsuoka 2001; Cho et al. 2001)

 Protoperidinium sp.
 Paleontological name: Stelladinium bifurcatum
 Distribution: East China Sea (Kim et al. 2010)
Remark: Stelladinium bifurcatum의 시스트 종명은 일부학술지에 등장하지만, 원 출저는 확인이 되지 않음.

 Protoperidinium sp.
 Paleontological name: Stelladinium robustum Zonneveld 1997 (Plate VI-9)
 Distribution: Busan harbor (Kim et al. 2005)

 Protoperidinium sp.
 Paleontological name: Trinovantedinium pallidifulvum Matsuoka 1987 (Plate VI-10)
 Distribution: Busan harbor (Kim et al. 2005), Gwangyang bay (Kim et al. 2003a), Gamak bay (Shin et al. 2010b; Shin et al. 2011), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Bokdol-bada (Shin et al. 2011), central South Sea (Cho et al. 2003)

Order GYMNODINIALES Lemmerman 1910
Family GYMNODINIACEA Lankester 1885
(Gymnodiniod Group)
Genus Polykrikos Kofoid et swezy 1921

 Polykrikos kofoidii Chatton 1914
 Paleontological name: cyst of Polykrikos kofoidii Chatton 1914 (Plate VI-11, 12)
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Kim 1991; Lee and Yoo 1991; Moon et al. 1993; Lee et al. 1998; Park and Shim 2011), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park and Yoon 2003; Park et al. 2004; Park and Shim 2011; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Kang et al. 1999; Pospelova and Kim 2010; Park and Shim 2011), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Lee and Matsuoka 1996; Cho et al. 2003; Park et al. 2005), Seamangeum coasts (Park et al. 2004; Park and Shim 2011), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2004; Kim et al. 2010)

 Polykrikos schwartzii Bütschli 1873
 Paleontological name: cyst of Polykrikos schwartzii Bütschli 1873 (Plate VI-13)
 Distribution: Busan harbor (Kim et al. 2005), Jinhae-Masan bay (Lee and Yoo 1991; Moon et al. 1993; Lee et al. 1998; Park and Shim 2011), Gwangyang bay (Kim et al. 2003a, b), Gamak bay (Lee et al. 1999; Park et al. 2004; Shin et al. 2010b; Shin et al. 2011; Shin et al. 2012), Yeoja bay (Shin et al. 2010a; Shin et al. 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Kang et al. 1999; Pospelova and Kim 2010), Bokdol-bada (Shin et al. 2011), western South Sea (Shin et al. 2007), central South Sea (Cho et al. 2003; Park et al. 2005), East China Sea (Cho and Matsuoka 2001; Cho et al. 2001; Cho et al. 2004; Kim et al. 2010)

 Polykrikos sp.
 Paleontological name: cyst of Polykrikos sp.
 Distribution: Jinhae-Masan bay (Kim 1991; Park and Shim 2011), Gamak bay (Shin et al. 2010b; Park and Shim 2011), Geoje coast (Shin et al. 2007), Tongyeong coasts (Park and Shim 2011), Seamangeum coasts (Park and Shim 2011), East China Sea (Cho et al. 2003; Kim et al. 2010)

3. 한국주변해역의 와편모조류 시스트 출현 특성

 한국 연안 해역의 와편모조류 시스트이 우점 출현종과 시스트의 세포밀도를 Table 4에 정리하였다. 한국 연안해역에서 Gymnodinium catenatum, Scrippsiella trochoidea, Lingulodinium polyedrum, Alexandrium spp. Cochlodinium polykrikoides 등과 같은 유해/유독 와편모조류는 특정 해역과 시기에 높은 세포밀도로 출현하여 적조를 형성하기도 하지만, 표층퇴적물에서 이들 시스트가 매우 높게출현한다. 1995년 이후 반복적으로 가을에 남해 고흥과 남면, 소리도 사이의 봇돌바다와 여수해만에서 적조를 발생시키는 C. polykrikoides에 대해 아직도 종자군(seed population)으로서 역할은 분명하지 않지만(Matsuoka et al. 2010), 시스트의 존재는 확실한 것으로 보고된다(Pospelova and Kim 2010). 그리고 1986년에 부산 및 1996년 거제도에서 각각 2명의 사망자를 배출한 남해안의 마비성 패류독화(Paralytic Shellfish Poisoning, PSP) 원인종으로 알려진 Alexandrium tamarense/catenella type (Han et al. 1992)은 전 세계에서 아시아 해역에 특히 높은 세포밀도를 보여(Marret and Zonneveld 2003; Zonneveld et al. 2013), 동중국해를 중심으로 중국연안(Wang et al. 2004)과 한국 연안 해역(Cho and Matsuoka 2001; Park et al. 2004; Shin et al. 2007)은 물론 일본 규슈서해안에 위치하는 아리아케만에서도 우점 출현하는 것이 보고된다(Shin, 2009). 국내에 유입되는 A. tamarense/catenella type의 기원은 분명하지 않지만, 이들 종의 분포적 특징(Zonneveld et al. 2013)으로부터 일본열도 남쪽에서 분기되어 동중국해와 서일본 그리고 한국 연안 해역에 영향을 미치는 쓰시마난류와 밀접한 관련성(Teague et al. 2003)을 보이는 것으로 추정되며, 진해-마산만 주변 해역은 A. tamarense 시스트의 집적과 발아, 그리고 유영세포의 성장에 좋은 환경 조건을 가지는 것으로 추정하고 있다(Shin et al. 2007). 또한 탄산염의 세포벽은 가지는 Scrippsiella trocoides 또한 한국 남해안에 우점 출현하고 있을 뿐만 아니라(Kang et al. 1999; Park et al. 2005, 2008), 해저 무산소와와 관련하여 해저환경의 pH 농도 변화에 따라 형태를 달리하는 것이 보고되고 있다(Shin et al. 2013).

 와편모조류 시스트 세포밀도는 적게는 개방된 외해에서 관찰이 되지 않은 정점도 있지만, 부영양화가 진행된 진해만(Lee et al. 1998)이나 통영 연안(Pospelova and Kim 2010)에서는 10,000 cysts/g dry 수준의 높은 세포밀도를 보여, 해역에 따른 변화 폭이 크게 나타난다(Table 4). 종별로는 Gonyaulacoid 그룹과 Protoperidinioid 그룹의 Spiniferites속과 Brigantedinium속의 종이 비교적 높은 밀도로 출현하나 해역에 따라 다소 다른 양상을 보인다. 일예로 가막만과 진해-마산만은 Briganntedinium속과 Protoperidinium속 같은 종속영양종이 많이 출현하는 반면에(Lee et al. 1998; Park and Yoon 2003; Shin 2009), 광양만이나 거제도 연안에서는 Spiniferites속의 S. bulloideus와 Alexandrium속 등의 독립영양종이 많이 출현하였다 (Kim et al. 2003a, b; Shin et al. 2007). 이와 같은 차이는 시스트가 퇴적되는 퇴적물의 입도조성(Dale 1983)과 와편모조류 유영세포 서식환경의 차이를 나타내는 것으로 판단된다(Smayda and Reynolds 2003). 즉 시스트는 미세한 세립질 퇴적물이 관찰되는 해역에서 높은 밀도를 보여, 펄 함량이 높은 남해 중앙부 해역보다 비교적 펄 함량이 낮은 진도, 완도가 위치한 남서연안에서 낮은 밀도로 관찰 된다(Shin et al. 2007). 시스트 분포는 이들 환경인자 이외에 수온, 광량, 염분, 영얌염류, 그리고 용존산소와 같은 물리∙화학적 환경인자 및 식물플랑크톤의 생물량과 세균의 활성도와 같은 생물학적 인자와도 밀접한 관련성을 보인다(Lee et al. 1998; Lee et al. 1999; Park and Yoon 2003; 서론 내용 참조).

Table 4. Abundance and dominant species of dinoflagellate cysts in the KCW

사 사

 이 논문은 2013년도 정부(교육과학기술부)의 재원으로 한국연구재단의 지원을 받아 수행된 기초연구사업(No. 2013005394)이 일부이다.

Reference

1.Alpermann TJ, B Beszteri, U John, U Tillmann and AD Cembella. 2009. Implications of life history transitions on the population genetic structure of the toxigenic marine dinoflagellate Alexandrium tamarense. Molecul. Ecol. 18:2122- 2133.
2.Anderson DM. 1984. The roles of dormant cysts in toxic dinoflagellate blooms and shellfish toxicity. pp.125-138. In Seafood Toxins (Ragelis E ed.). American Chem. Soc. Symp. Series. Washington, DC.
3.Anderson DM and BA Keafer. 1987. An endogenous annual clock in the toxic marine dinoflagellate Gonyaulax tamarensis. Nature 325:616-617.
4.Anderson DM, CD Taylor and EV Armbrust. 1987. The effects of darkness and anaerobiosis on dinoflagellate cyst germina-tion. Limnol. Oceanogr. 32:340-351.
5.Anderson DM and K Rengefors. 2006. Community assembly and seasonal succession of marine dinoflagellates in a temperate estuary: The importance of life cycle events. Limnol. Oceanogr. 51:860-873.
6.Anderson DM, SW Chisholm and CJ Watras. 1983. Importance of life cycle events in population dynamics of Gonyaulax tamarensis. Mar. Biol. 76:179-189.
7.Anderson DM, TJ Alpermann, AD Cembella, Y Collos, E Masseret and M Montresor. 2012. The globally distributed genus Alexandrium: multifaceted roles in marine ecosystems and impacts on human health. Harmful Algae 14:10-35.
8.Andersoen JT. 1998. The effect of seasonal variability on the germination and vertical transport of a cyst forming dinoflagellate, Gyrodinium sp., in the Chesapeake Bay. Ecol. Model. 112:85-109.
9.Baula IU, RV Azanza, Y Fukuyo and FP Siringan. 2011. Dinoflagellate cyst composition, abundance and horizontal distribution in Bolinao, Pangasinan, Northern Philippines. Harmful Algae 11:33-44.
10.Bazzoni AM, M Bastianini, M Montresor and C Totti. 2006. Annual cycle of phytoplankton assemblages and first studies of vertical fluxes of dinoflagellate cysts in the North-western Adriatic Sea. Biol. Mar. Mediterranea 13:940-942.
11.Bibby BT and JD Dodge. 1972. The encystment of a freshwater dinoflagellate: a light and electron-microscopical study. British Phycol. J. 7:85-100.
12.Bolch CJS and MF de Salas. 2007. A review of the molecular evidence for ballast water introduction of the toxic dinoflagellates Gymnodinium catenatum and the Alexandrium "tamarensis complex" to Australasia. Harmful Algae 6: 465-485.
13.Bonnet S, Ade Vernal, R Gersonde and L Lembke-Jene. 2012. Modern distribution of dinocysts from the North Pacific Ocean (37-64˚N, 144˚E-148˚W) in relation to hydrographic conditions, sea-ice and productivity. Mar. Micropaleontol. 84/85:87-113.
14.Bravo I, RI Figueroa, E Garcés, S Fraga and A Massanet. 2010. The intricacies of dinoflagellate pellicle cysts: the example of Alexandrium minutum cysts from a bloom-recurrent area (Bay of Baiona, NW Spain). Deep-Sea Res. II, 57:166-174.
15.Candel MS, T Radi, Ade Vernal and G Bujalesky. 2012. Distribution of dinoflagellate cysts and other aquatic palyno-morphs in surface sediments from the Beagle Channel, Southern Argentina. Mar. Micropaleontol. 96/97:1-12.
16.Cho HJ, CH Kim, CH Moon and K Matsuoka. 2003. Dinoflagellate cysts in recent sediments from the southern coastal waters of Korea. Bot. Mar. 46:332-337.
17.Cho HJ, JB Lee and CH Kim. 2004. Dinoflagellate cyst distribution in the surface sediments from the East China Sea around Jeju Island. Korean J. Environ. Biol. 22:192-199. (in Korean)
18.Cho HJ and K Matsuoka. 2001. Distribution of dinoflagellate cysts in surface sediments from the Yellow Sea and East China Sea. Mar. Micropaleont. 42:103-123.
19.Cho HJ and K Matsuoka, 2003. Analysis of marine environmental changes based on dinoflagellate cyst biostratigraphy in the East China Sea. Bull. Mar. Envion. Res. Inst. 27:9-13.
20.Cho HJ, K Matsuoka, JB Lee and CH Moon. 2001. Dinoflagellate cyst assemblages in the surface sediments from the northern East China Sea. J. Fish. Sci. Tech. 4:120-129.
21.Corrales RA and Jl Maclean. 1995. Impact of harmful algae on sea farming in the Asia-Pacific areas. J. Appl. Phycol. 7: 151-162.
22.Cox AM, DH Shull and RA Horner. 2008. Profiles of Alexandrium catenella cysts in Puget Sound sediments and the relationship to paralytic shellfish poisoning events. Harmful Algae 7:379-388.
23.Dai X, D Lu and C Wang. 2012. Analysis of the population structure of dinoflagellate cysts in the sediment of ballast tank of four cargo boats at Zhoushan Port. J. Mar. Sci. 30: 10-18.
24.Dale B. 1977. Cysts of the toxic red-tide dinoflagellate Gonyaulax excavata (Braarud) Balech from Oslo fjorden, Norway. Sarsia 63:29-34.
25.Dale B. 1983. Dinoflagellate resting cyst. pp.69-144. In Survival Strategies of the Algae (Fryxell GA ed.), Cambridge University Press, Cambridge.
26.Dale B. 2000. Dinoflagellate cysts as indicators of cultural eutrophication and industrial pollution in coastal sediments. Environ. Micropaleontol. (Topics in Geobiology) 15:305- 321.
27.Dale B. 2001a. Marine dinoflagellate cysts as indicators of eutrophication and industrial pollution: a discussion. Sci. Total Environ. 264:235-240.
28.Dale B. 2001b. The sedimentary record of dinoflagellate cysts: looking back into the future of phytoplankton blooms. Sci. Mar. 65:257-272.
29.Dale B. 2009. Eutrophication signals in the sedimentary record of dinoflagellate cysts in coastal waters. J. Sea Res. 61:103- 113.
30.Dale B, CM Yentsch and JW Hurst. 1978. Toxicity in resting cyst of red-tide dinoflagellate Gonyaulax excavata from deeper water coastal sediments. Science 201:1223-1225.
31.Dale B, TA Thorsen and A Fjellsa. 1999. Dinoflagellate cysts as indicators of cultural eutrophication in the Oslofjord, Norway. Est. Coast. Shelf Sci. 48:371-382.
32.de Vernal A and A Rochon. 2011. Dinocysts as tracers of sea-surface conditions and sea-ice cover in polar and subpolar environments, IOP Conference Series. Earth Environ. Sci. 14:1-12 (012007).
33.Devillers R and A de Vernal. 2000. Distribution of dinoflagellate cysts in surface sediments of the northern North Atlantic in relation to nutrient content and productivity in surface waters. Mar. Geol. 166:103-124.
34.Elleraard M, DM Kulis and DM Anderson. 1998. Cyst of Danish Gymnodinium nolleri Ellegaard et Moestrum sp. ined. (Dinophyceae): studies on encystment, excystment and toxicity. J. Plankton Res. 20:1743-1755.
35.Elshanawany R, K Zonneveld, M Ibrahim and SEA Kholief. 2010. Distribution patterns of recent organic-walled dinoflagellate cysts in relation to environmental parameters in the Mediterranean Sea. Palynol. 34:233-260.
36.Fahnenstiel G, Y Hong, D Millie, M Doblin, T Johengen and D Reid. 2009. Marine dinoflagellate cysts in the ballast tank sediments of ships entering the Laurentian Great Lakes. Verhandlungen Intern. Verein Limnol. 30:1035-1038.
37.Fensome RA, FJR Taylor, G Norris, WAS Sarjeant, DI Wharton and GL Williams. 1993. A Classification of Modern and Fossil Dinoflagellates. Micropaleontology Special Publication 7. Sheridan Press, New Hampshire, 351pp.
38.Figueroa RI, JA Vazquez, A Massanet, MA Murado and I Bravo. 2011. Interactive effects of salinity and temperature on planozygote and cyst formation of Alexandrium minutum (Dinophyceae) in culture. J. Phycol. 47:13-24.
39.Grøsfjeld K, R Harland and J Howe. 2009. Dinoflagellate cyst assemblages inshore and offshore Svalbard reflecting their modern hydrography and climate. Norwegian J. Geol. 89: 121-134.
40.Gu H, Q Fang, J Sun, D Lan, F Cai and ZY Gao. 2003. Dinoflagellate cysts in recent marine sediment front Guangxi, China. Acta Oceanol. Sinica 22:407-419.
41.Guiot J and A de Vernal. 2011. Is spatial autocorrelation introducing biases in the apparent accuray of paleoclimatic reconstructions? Quatern. Sci. Rev. 30:1965-1972.
42.Hallegraeff GM and CJ Bolch. 1992. Transport of diatom and dinoflagellate resting spores in ships' ballast water: implications for plankton biogeography and aquaculture. J. Plankton Res. 14:1067-1084.
43.Hallegraeff GM, CJ Bolch, J Bryan and B Koerbin. 1990. Microalgal sporea in ships' ballast water: A danger to aquaculture. pp.475-480. In Toxic Marine Phytoplankton (Graneli E et al. eds.). Elsevier, New York.
44.Hallegraeff GM, SO Stanly, CJ Bolch and SI Blackburn. 1989. Gymnodinium catenatum blooms and shellfish toxicity in southern Tasmania. pp.77-80. In Red Tides: Biology, Environmental Science, and Toxicology (Okaichi T, DM Anderson and K Nemoto eds). Elsevies, New York.
45.Hamer JP, IAN Lucas and TA McCollin. 2001. Harmful dinoflagellate resting cysts in ships' ballast tank sediments: potential for introduction into English and Welsh waters. Phycologia 40:246-255.
46.Han MS, JK Jeon and YO Kim. 1992. Occurrence of dinoflagellate Alexandrium tamarense, a causative organism of paralytic shellfish poisoning in Chinhae Bay. Korea. J. Plankton Res. 14:1581-1592.
47.Harper FM, EA Hatfield and RJ Thompson. 2002. Recirculation of dinoflagellate cysts by the mussel, Mytilus edulis L., at an aquaculture site contaminated by Alexandrium fundyense (Lebour) Balech. J. Shellfish Res. 21:471-477.
48.Head MJ. 1996. Modern dinoflagellate cysts and their biological affinies. pp.1197-1248. In Palynology: Principles and Applications, vol. 3 (Jansonius J and DC McGregor eds.), American Ass. Stratigr. Palynol. Found. Texas.
49.Heiskanen AS. 1993. Mass encystment and sinking of dinoflagellates during a spring bloom. Mar. Biol. 116:161-167.
50.Hong Y, GL Fahnenstiel and R Stone. 2003. Dinoflagellate cysts in NOBOB ballast tank sediment: potential risk of invasion to the Great Lakes. Intern. Ass. Great Lakes Res. [URL: http://iaglr.org/].
51.Hoppenrath M and BS Leander. 2007. Character evolution in Polikrikoid dinoflagellates. J. Phycol. 43:366-377.
52.Hoppenrath M, M Elbrachter and G Drebes. 2009. Marine phytoplankton. selected microphytoplankton species from the North Sea around Helgoland and Sylt. E. Schweizerbart'sche Verlagsbuchhandlung (Nägele u. Obermiller). Stuttgart, 114pp.
53.Hwang CH, S Heo and CH Kim. 2009. Horizontal distribution of dinoflagellate resting cysts in sediments from the southeastern Yellow Sea. J. Korean Fish. Soc. 42:68-72. (in Korean)
54.Ichikawa S, Y Wakao and Y Fukuyo. 1992. Extermination efficacy of hydrogen peroxide against cysts of red tide and toxic dinoflagellates, and its adaptability to ballast water of cargo ships. Nippon Suisan Gakkaishi 58:2229-2233. (in Japanese)
55.Joyce LB and C Pitcher. 2004. Encystment of Zygabikodinium lenticulatum (Dinophyceae) during a summer bloom of dinoflagellates in the southern Benguela upwelling system. Est. Coast. Shelf Sci. 59:1-11.
56.Joyce LB, GC Pitcher and PA Cook. 2002. Dinoflagellate cysts and the initiation of harmful blooms on the west coast of South Africa. Southern African Network for Coastal and Oceanographic Research (SANCOR), Benguela Environment Fisheries Interaction and Training Programme BENEFIT).
57.Kang YJ, CH Moon and HJ Cho. 2008. Comparison of panning and sodium polytungstate methods for separating dinoflagellate cysts. J. Korean Fish. Soc. 41:228-231. (in Korean)
58.Kang YJ, TH Ko, JA Lee, JB Lee and IK Chung. 1999. The community dynamics of phytoplankton and distribution of dinoflagellate cysts in Tongyoung Bay, Korea. Algae 14:43- 54.
59.Kim CH. 1994. Germinability of resting cysts associated with occurrence of toxic dinoflagellate Alexandrium species. J. Aquacul. 7:251-264.
60.Kim CH. 1995. Paralytic shellfish toxin profiles of the dinoflagellate Alexandrium species isolated from benthic cysts in Jinhae bay, Korea. J. Korean Fish. Soc. 28:364-372.
61.Kim CH. 2009. Life cycle of the harmful dinoflagellate Cochlodinium polykrikoides. Bull. Plankton Soc. Japan 56:31.
62.Kim CH, HJ Cho, JB Shin, CH Moon and K Matsuoka. 2002. Regeneration from hyaline cysts of Cochlodinium polykrikoides (Gymnodiniales, Dinophyceae). a red tide organism along the Korean coast. Phycol. 41:667-669.
63.Kim CJ, HG Kim, CH Kim and HM Oh. 2007. Life cycle of the ichthyotoxic dinoflagellate Cochlodinium polykrikoides in Korean coastal waters. Harmful Algae 6:104-111.
64.Kim HG. 1991. The distribution and role of benthic cysts on the dinoflagellate bloom in Chinhae bay. pp.99-120.
65.In Recent Approaches on Red Tides (Park JS and HK Kim eds.). National Fish. Res. & Dev. Agency, Busan. Kim HG, JS Park and SG Lee. 1990. Coastal algal blooms caused by the cyst-forming dinoflagellates. J. Korean Fish. Soc. 23:468-474.
66.Kim HJ, CH Moon and HJ Cho. 2005. Spacial-temporal characteristics of dinoflagellate cyst distribution in sediments of Busan harbor. J. Korean Soc. Oceanogr. (The Sea) 10:196- 203. (in Korean)
67.Kim SY, CH Moon and HJ Cho. 2003a. Relationship between dinoflagellate cyst distribution in surface sediments and phytoplankton assemblages from Gwangyang bay, a souther coastal area of Korea. J. Korean Soc. Oceanogr. (The Sea) 8:111-120. (in Korean)
68.Kim SY, CH Moon and HJ Cho. 2003b. Vertical distribution of dinoflagellate cysts in sediments from Gwangyang bay, Korea. J. Korean Fish. Soc. 36:290-297. (in Korean)
69.Kim SY, CH Moon, HJ Cho and DI Lim. 2009. Dinoflagellate cysts in coastal sediments as indicators of eutrophication: A case of Kwangyang bay, South Sea of Korea. Est. Coasts 32:1225-1233.
70.Kim SY, DI Lim and SR Kang. 2010, Dinoflagellate cyst assemblages in a sediment core from the northern East China Sea and their implications. J. Geol. Soc. Korea 46:221-231.
71.Kim YO, MH Park and MS Han. 2002. Role of cyst germination in the bloom iniation og Alexandrium tamarense (Dinophyceae) in Masan Bay. Aqua. Microbial. Ecol. 29:279-286.
72.Kim YO and MS Han. 2000. Seasonal relationships between cyst germination and vegetative population of Scrippsiella trochoidea (Dinophyceae). Mar. Ecol. Pro. Ser. 204:111- 118.
73.Kita T, Y Fukuyo, H Tokuda and R Hirano. 1993. Sexual reproduction of Alexandrium hiranoi (Dinophyceae). Bull. Planton Soc. Japan 39:79-85.
74.Kremp A, DH Shull and DM Anderson. 2003. Effects of deposit- feeder gut passage and fecal pellet encapsulation on germination of dinoflagellate resting cysts. Mar. Ecol. Prog. Ser. 263:65-73.
75.Kremp A and DM Anderson. 2000. Factors regulating germination of resting cysts of the spring bloom dinoflagellate Scrippsiella hangoei from the northern Baltic Sea. J. Plankton Res. 22:1311-1327.
76.Kunz-Pirrung M, J Matthiessen and A de Vernal. 2001. Late Holocene dinoflagellate cysts as indicators for short-term climate variability in the eastern Laptev Sea (Arctic Ocean), J. Quatern. Sci. 16:711-716.
77.Lan D, Q Fang, H Gu and C Li. 2004. Resting cysts of harmful and toxic dinoflagellates and potential damage of Alexandrium tamarense from Xiamen Bay sediments. J. Oceanogr. Taiwan Strait 23:453-457.
78.Lee JB, DY Kim and JA Lee. 1998. Community dynamic and distribution of dinoflagellates and their cysts in Masan- Chinhae Bay, Korea. J. Fish. Sci. Tech. 1:283-292.
79.Lee JB and K Matsuoka. 1996. Dinoflagellate cysts in surface sediments of southern Korean waters. p.173-176. In Harmful and Toxic Algal Blooms (Yasumoto T, Y Oshima and Y Fukuyo (eds.). IOC of Unesco, Paris.
80.Lee JB and KI Yoo. 1991. Distribution of dinoflagellate cysts in Masan Bay, Korea. J. Oceanogr. Soc. Korea 26:304-312.
81.Lee MH, JB Lee, JA Lee and JG Park. 1999. Community structure of flagellates and dynamics of resting cysts in Kamak bay, Korea. Algae 14:255-266. (in Korean)
82.Marret F and KAF Zonneveld. 2003. Atlas of modern organicwalled dinoflagellate cyst distribution. Rev. Palaeobot. Palynol. 125:1-200.
83.Matrai P, B Thompson and M Keller. 2005. Circannual excystment of resting cysts of Alexandrium spp. from eastern Gulf of Maine populations. Deep-Sea Res. II 52:2560-2568.
84.Matsuoka K. 1999. Eutrophication process recorded in dinoflagellate cyst assemblages-a case of Yokohama Port, Tokyo Bay, Japan. Sci. Total Environ. 231:17-35.
85.Matsuoka K. 2001. Further evidence for a marine dinoflagellate cyst as an indicator of eutrophication in Yokohama Port, Tokyo Bay, Japan. Comments on a discussion by B. Dale. Sci. Total Environ. 264:221-233.
86.Matsuoka K. 2011. Dinoflagellate cyst as a bio-signal for eutrophication. Bull. Plankton Soc. Japan 58:55-59. (in Japanese).
87.Matsuoka K, A Mizuno, M Iwataki, Y Takano, T Yamatogi, YH Yoon and JB Lee. 2010. Seed populations of a harmful unarmored dinoglagellate Cochlodinium polykrikoides Margalef in the East China Sea. Harmful Algae 9:548-556.
88.Matsuoka K and Y Fukuyo. 2000. Technical guide for modern diniflagellate cyst study. WESTPAC-HAB/WESTPAC/IOC, 29pp., 24 figures, 21 plates.
89.Matsuyama Y. 2012. Impacts of the harmful dinoflagellate Heterocapsa circularisquama bloom on shellfish aquaculture in Japan and some experimental studies on invertebrates. Harmful Algae 14:144-155.
90.Moon SK, GY Ryoo, JN Lee and CK Hong. 1993. Classification of dinoflagellate resting cysts in Chinhae bay. Korea. Bull. Kyungsung Univ. 12:129-148. (in Korean)
91.Morey-Gains G and RH Ruse. 1980. Encystment and reproduction of the predatory dinoflagellate, Polykrikos kofoidii Chatton (Gymnodiniales). Phycologia 19:230-232.
92.Naidu PD, JS Patil, DD Narale and AC Anil. 2012. A first look at the dinoflagellate cysts abundance in the Bay of Bengal: implications on Late Quaternary productivity and climate change. Current Sci. 102:495-499.
93.Nehring S. 1997. Dinoflagellate resting cysts in recent sediments of the western Baltic as indicators for the occurrence of 'nonindigenous' species in the water column. pp.79-85. In Proceedings of the 13th Symposium of the Baltic Marine Biologists (Andrusaitis A ed.). Institute of Aquatic Ecology, University of Latvia.
94.Odebrecht C. 1982. The production of resting cysts by planktonic organisms as a survival strategy in Harrington Sound, Bermuda. Atlantica 5:87.
95.Park JS and YH Yoon. 2003. Marine environmental characteristics by distribution of dinoflagellate cysts in the Southwestern coastal waters of Korea 1. Spatio-temporal distribution of dinoflagellate cysts in Gamak Bay. J. Korean Fish. Soc. 36:151-156. (in Korean)
96.Park JS, YH Yoon and IH Noh. 2004. Estimation of the variation of marine environment by the distribution of organic matter and dinoflagellate cysts in the vertical sediments in Gamak Bay, Korea. J. Korean Soc. Mar. Environ. Eng. 7: 164-173. (in Korean)
97.Park JS, YH Yoon, IH Noh and HY Soh. 2005. A study of organic matter and dinoflagellate cyst on surface sediments in the central parts of South Sea, Korea. Korean J. Environl. Biol. 23:163-172. (in Korean)
98.Park JS, YH Yoon, IH Noh, HY Soh and HH Shin. 2008. The marine environment and dinoflagellates cysts in the Southwestern Sea of Korea. Algae 23:135-140. (in Korean)
99.Park JY and JH Shim. 2011. Dinoflagellate resting cysts from surface sediments of Korean coastal waters. pp.132-150. In The plankton ecology of Korean coastal waters (Choi, C.K. eds). Donghwa Press, Seoul. (in Korean)
100.Park KH, KY Kim, CH Kim and HG Kim. 2004. Spatio-temporal distribution of dinoflagellate resting cysts at the Saemangeum area. J. Korean Fish. Soc. 37:202-208.
101.Park MH, YO Kim, SH Cho and MS Han. 2004. Effects of environmental conditions on germination of Alexandrium tamarense cysts from Masan bay, Korea. Korean J. Environ. Biol. 22:200-205. (in Korean)
102.Park TG, YT Park and HM Bae. 2009. Life cycle of heterotrophic dinoflagellate Cryptoperidiniopsis brodyi (Dinophyceae). J. Environ. Sci. 18:9-14. (in Korean)
103.Patil JG, RM Gunasekera, BE Deagle, NJ Bax and SI Blackburn. 2005. Development and evaluation of a PCR based assay for detection of the toxic dinoflagellate, Gymnodinium catenatum (Graham) in ballast water and environmental samples. Biol. Invasions 7:983-994.
104.Persson A. 2000. Possible predation of cysts-a gap in the knowledge of dinoflagellate ecology? J. Plankton Res. 22:803- 809.
105.Persson A and BC Smith. 2009a. Grazing on a natural assemblage of ciliate and dinoflagellate cysts by the eastern oyster Crassostrea virginica. Aquatic Biol. 6:227-233.
106.Persson A and BC Smith. 2009b. Consumption of Scrippsiella lachrymosa resting cysts by the eastern oyster (Crassostrea virginica). J. Shellfish Res. 28:221-225.
107.Pertola S, MA Faust and H Kuosa. 2006. Survey on germination and species composition of dinoflagellates from ballast tanks and recent sediments in ports on the South Coast of Finland, North-Eastern Baltic Sea. Mar. Poll. Bull. 52:900- 911.
108.Pospelova V and SJ Kim. 2010. Dinoflagellate cysts in recent estuarine sediments from aquaculture sites of southern South Korea. Mar. Micropaleontol. 76:37-51.
109.Pospelova V, TF Pedersen and A de Vernal. 2006. Dinoflagellate cysts as indicators of climatic and oceanographic changes during the past 40 kyr in the Santa Barbara Basin, southern California. Paleoceanogr. 21:1-16.
110.Price AM, KNM Mertens, V Pospelova, TF Pedersen and RS Ganeshram. 2013. Late Quaternary climatic and oceanographic changes in the Northeast Pacific as recorded by dinoflagellate cysts from Guaymas Basin, Gulf of California (Mexico). Paleoceanogr. DOI:10.1002/palo.20019.
111.Price AM and V Pospelova. 2011. High-resolution sediment trap study of organic-walled dinoflagellate cyst production and biogenic silica flux in Saanich Inlet (BC, Canada). Mar. Micropaleontol. 80:18-43.
112.Sarjenat WAS, T Lacalli and G Gains, 1987. The cysts and skeletal elements of dinoflagellates: speculations on the ecological causes for their morphology and development. Micropaleontol. 33:1-36.
113.Satta CT, S Anglès, A Lugliè, J Guillén, N Sechi, J Camp and E Garcés. 2013. Studies on dinoflagellate cyst assemblages in two estuarine Mediterranean bays: A useful tool for the discovery and mapping of harmful algal species. Harmful Algae 24:65-79.
114.Sgrosso S, F Esposito and M Montresor. 2001. Temperature and daylength regulate encystment in calcareous cyst-forming dinoflagellates. Mar. Ecol. Pro. Ser. 211:77-87.
115.Shin HH, JS Park, YO Kim, SH Baek, DI Lim and YH Yoon. 2012. Dinoflagellate cyst production and flux in Gamak Bay, Korea: A sediment trap study. Mar. Micropaleontol. 94/95:72-79.
116.Shin HH, K Matsuoka, YH Yoon, YO Kim. 2010a. Response of dinoflagellate cyst assemblages to salinity changes in Yeoja Bay, Korea. Mar. Micropaleontol. 77:15-24.
117.Shin HH, K Mizushima, SJ Oh, JS Park, IH Noh, M Iwataki, K Matsuoka and YH Yoon. 2010b. Reconstruction of historical nutrient levels in Korean and Japanese coastal areas based on dinoflagellate cyst assemblages. Mar. Poll. Bull. 60:1243-1258.
118.Shin HH, SW Jung, MC Jang and YO Kim, 2013. Effect of pH on the morphology and viability of Scrippsiella trochoidea cysts in the hypoxia zone of eutrophied area. Harmful Algae. 28:37-46.
119.Shin HH, YH Yoon, H Kawami, M Iwataki and K Matsuoka. 2008. The first appearance of toxic dinoflagellate Alexandrium tamarense (Gonyaulacales, Dinophyceae) responsible for the PSP contaminations in Gamak Bay, Korea. Algae 23:251-255.
120.Shin HH, YH Yoon and JS Park. 2007. Marine Environmental Characteristics on the Dinoflagellate cysts Distribution in Surface Sediments in the Southwest Sea, Korea. Korean J. Environ. Biol. 25:205-214. (in Korean)
121.Shin HH, YH Yoon and K Matsuoka. 2007. Modern dinoflagellate cyst distribution off the eastern part of Geoje Island, Korea. Ocean Sci. J. 42:31-39.
122.Shin HH, YH Yoon, YO Kim and K Matsuoka. 2011. Dinoflagellate cysts in surface sediments from Southern Coast of Korea. Est. Coasts 34:712-725.
123.Smayda TJ. 2008. Complexity in the eutrophication - harmful algal bloom relationship, with comment on the importance of grazing. Harmful Algae 8:140-151.
124.Smayda TJ and CS Reynolds. 2003. Strategies of marine dinoflagellate survival and some rules of assembly. J. Sea Res. 49:95-106.
125.Solignac S, A de Vernal and J Giraudeau. 2008. Comparison of coccolith and dinocyst assemblages in the northern North Atlantic: How well do they relate with surface hydrography? Mar. Micropaleontol. 68:115-135.
126.Sorrel P, SM Popescu, MJ Head, JP Suc, S Klotz and H Oberhänsli. 2006. Hydrographic development of the Aral Sea during the last 2000 years based on a quantitative analysis of dinoflagellate cysts. Palaeogeo. Palaeoclim. Palaeoecol. 234:304-327.
127.Steidinger KA. 1975. Basic factors influencing red tides. pp. 249-255. In Proc. 1st Conf. on Toxic Dinoflagellate Blooms (Locicero VR ed.). Mass. Sci. Technol. Found, Wakefield.
128.Takeuchi T. 1985. Ecology in west coast of Kii Peninsula. pp. 98-108. In Toxic Dinoflagellate -Implication in Shellfish Poisoning- (Fukuyo Y ed.). Koseisha-Koseikaku, Tokyo. (in Japanese)
129.Tang YZ and CJ Gobler. 2012, The toxic dinoflagellate Cochlodinium polykrikoides (Dinophyceae) produces resting cysts. Harmful Algae 20:71-80.
130.Taylor FJR. 1987. The Biology of Dinoflagellates. Blackwell Sci. Publ. London, 785pp.
131.Taylor FJR, M Hoppenrath and JF Saldarriaga. 2008. Dinoflagellate diversity and distribution. Biodiv. Conserv. 17:407-418.
132.Teague WJ, GA Jacobs, DS Ko, TY Tang, KI Chang and MS Suk. 2003. Connectivity of the Taiwan, Cheju, and Korea Straits. Conti. Shelf Res. 23:63-67.
133.Tyler MA, DM Coasts and DM Anderson. 1982. Encystment in a dynamic environment: deposition of dinoflagellate cysts by a frontal convergence. Mar. Ecol. Prog. Ser. 7:163-178.
134.Uchida T, Y Matsuyama, M Yamaguchi and T Honjo. 1996. The life cycle of Gyrodinium instriatum (Dinophyceae) in culture. Phycol. Res. 44:119-123.
135.Usup G, A Ahmad, K Matsuoka, PT Lim and CP Leaw. 2012. Biology, ecology and bloom dynamics of the toxic marine dinoflagellate Pyrodinium bahamense. Harmful Algae 14: 301-312.
136.von Stosch HA. 1973. Observation on vegetative reproduction and sexual life cycles of two freshwater dinoflagellates, Gymnodinium pseudopalstre Schiller and Woloszynskia apiculata sp. nov. British Phycol. J. 8:105-134.
137.Walker LM. 1984. Life histories, dispersal, and survival in marine prvival in dinoflagellates. pp.19-34. In Steidinger KA and LM Walker (eds), Marine Survival Life Cycle Strategies, CRCds)ess, Inc, Florida.
138.Walker LM and KA Steidinger. 1979. Sexual reproduction in the toxic dinoflagellate Gonyaulax monilata. J. Phycol. 15: 312-315.
139.Wall D and B Dale. 1968. Modern dinoflagellate cysts and evolution of the Peridiniales. Microplaleontol. 14:265-304.
140.Wall D, B Dale, GP Lohmann and WK Smith. 1977. The environmental and climatic distribution of dinoflagellate cysts in modern marine sediments from regions in the North and South Atlantic Oceans and adjacent seas. Mar. Micropaleontol. 2:121-200.
141.Wang Z, K Matsuoka, Y Qi and J Chen. 2004. Dinoflagellate cysts in recent sediments from Chinese coastal waters. Mar. Ecol. 25:289-311.
142.Wang Z, Y Qi and YF Yang. 2007. Cyst formation: an important mechanism for the termination of Scrippsiella trochoidea (Dinophyceae) bloom. J. Plankton Res. 29:209-218.
143.Yoo JS and HW Shin. 2004. Effects of basic oxygen furnace slag and inorganic nutrients on the germination of resting cysts of two toxic dinoflagellates. J. Environ. Biol. 25:147- 150.
144.Zonneveld KAF, F Marret, GJM Versteegh, K Bogus, S Bonnet, I Bouimetarhan, E Crouch, A de Vernal, R Elshanawany, L Edwards, O Esper, S Forke, K Grøsfjeld, M Henry, U Holzwarthc, JF Kielt, SY Kim, S Ladouceur, D Ledu, L Chen, A Limoges, L Londeix, SH Lu, MS Mahmoud, G Marino, K Matsouka, J Matthiessen, DC Mildenhal, P Mudie, HL Neil, V Pospelova, Y Qi, T Radi, T Richerol, A Rochon, F Sangiorgi, S Solignac, JL Turon, T Verleye, Y Wang, Z Wang and M Young. 2013. Atlas of modern dinoflagellate cyst distribution based on 2405 datapoints, Rev. Palaeobot. Palynol. in press