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Phylogeny of the Family Ophioglossaceae with Special Emphasis on Genus Mankyua

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Phylogeny of the Family Ophioglossaceae with Special Emphasis on Genus Mankyua Korean J. Pl. Taxon. (2009) Vol. 39 No. 3, pp. 135-142 제주고사리삼을 중심으로한 고사리삼과 식물의 계통 선병윤*·백태규·김영동1·김찬수2 전북대학교 생물과학부, 1한림대학교 생명과학과, 2국립산림과학원 난대산림연구소 Phylogeny of the family Ophioglossaceae with special emphasis on genus Mankyua Byung-Yun Sun*, Tae Gyu Baek, Young-Dong Kim1 and Chan Soo Kim2 Division of Biological Sciences, Chonbuk National University, Jeonju 561-756, Korea 1Division of Biological Sciences, Hallym University, 39 Hallymdaehak-gil, Chuncheon 200-702, Korea 2Warm-temperate Forest Research Center, Korea Forest Research Institute, Sanghyo-Dong, Seogwipo 697-050, Korea 적요: 엽록체 rbcL gene의 염기서열과 포자의 형태를 바탕으로 고사리삼과 식물의 계통과 제주고사리삼속 의 계통학적 위치를 추정하였다. 엽록체 DNA rbcL 염기서열의 분석 결과 고사리삼과는 뚜렷하게 고사리삼계 보 (Botrychioid lineage)와 나도고사리삼계보 (Ophioglossoid lineage)의 두 군으로 구분되었다. 고사리삼계보에 있어 제주고사리삼속(Mankyua)은 Helmintostachys속과 함께 분계의 기저에서 고사리삼속(Botrychium)의 자매분 류군으로 분지하였으나, 이들 고사리삼속(Botrychium), 제주고사리삼속(Mankyua), 및 Helmintostachys속 사이의 계통적 유연관계는 결정되지 못했다. 고사리삼속의 경우 Sceptridium아속과 Botrychium아속이 뚜렷한 단계통을 형성하였으나, Botrypus아속에 속한 분류군들은 고사리삼속의 기저에서 다른 고사리삼속 분류군의 자매분류군 으로 분지하면서 단계통이 아닌 것으로 밝혀졌다. 나도고사리삼계보를 형성하는 나도고사리삼속(Ophioglossum) 의 경우, Ophioglossum아속은 단계통을 형성하였으며, Ophioglossum아속에 대해 Cheiroglossa아속 및 Ophioderma 아속이 차례로 자매분류군으로 분지하였다. 염기서열 계통수, 포자의 형태 및 외부형태의 고찰에 있어 제주고 사리삼속은 Helminthostachys와 유사하지만, 계통수에서 patristic distance 뿐 아니라 포자소엽의 형태가 뚜렷이 구분되어 이들 2속은 독립된 속으로 판단된다. 주요어: 고사리삼과, 제주고사리삼속, 고사리삼속, 나도고사리삼속, Helminthostachys속, 분자계통, 포자 형태 ABSTRACT: Phylogeny of the family Ophioglossaceae and a phylogenetic position of Mankyua were estimated through analyses of chloroplast rbcL gene sequences and spore morphology. Sequence analysis of the rbcL gene clearly indicated that there are two major lineages in the family Ophioglossaceae: Botrychioid lineage and Ophioglossoid lineage. The Botrichioid lineage is composed of three distinct clades: Botrychium, Helminthostachys and Mankyua, where Helminthostachys and Mankyua were placed as sister groups to the Botrychium. Within the genus Botrychium, subgenera Septridium and Botrychium were monophyletic, while taxa of subgen. Botrypus branched as sister of the two, successively, thus making a non-monophyletic group. Ophioglossum formed the Ophioglossoied lineage, where the subgen. Ophioglossum is monophyletic, while subgen. Cheiroglossa and Ophoderma formed a sister relationship with subgen. Ophioglossum. In terms of external morphology and spores, Mankyua is most similar to Helminthostachys, however, patristic distance in the cladogram and trophophore characteristics of the two genera are distinct. Therefore, Mankyua is a well defined genus within the family in terms of morphology as well as molecular phylogeny which places it in basal position of the Botrychioid lineage on the gene tree. Keywords: Ophioglossaceae, Mankyua, Botrychium, Ophioglossum, Helminthostachys, molecular phylog- eny, spore morphology *Author for correspondence: [email protected] 135 136 선병윤·백태규·김영동·김찬수 비종자관속식물 (seedless vascular plant) 또는 관속포자 특징을 지닌다. 나도고사리삼속은 영양엽편이 단엽으로 식물 (spore bearing vascular plants)은 최근까지 솔잎난식물 전연이며 망상맥을 가지는 특징을 지닌다. 또한 포자낭수 문 (Psilophyta), 석송식물문 (Lycophyta), 속새식물문(Equi- 는 다육질로서 포자낭이 두 줄로 포자낭수 주변부에 매몰 setophyta) 및 양치식물문 (Filicophyta)의 4개의 문으로 인 되어 있고 지하경은 직립하며, 뿌리 끝에 흔히 주아가 달 식되어 왔다 (Gilford and Foster, 1989). 한편, 분자계통학적 려 영양번식을 한다. 제주고사리삼속의 경우, 영양엽편은 연구에서는 관속식물을 석송식물군 (또는 소엽식물군: Helminthostachys와 유사한 3출 복엽이지만, 포자엽편은 Lycophyte)과 진정엽식물군 (또는 대엽식물군: Euphyllophyte) 나도고사리삼속과 같이 다육질의 포자낭수에 포자낭이 으로 구분하고, 후자에 종자식물군(Spermatophyte)과 양치 두 줄로 포자낭수 주변부에 매몰되어 있으며, 지하경은 식물군 (Monilophyte)을 두어, 비종자관속식물을 단계통 직립하고 뿌리 끝에 흔히 주아가 달려 영양번식을 한다. 군으로 구분한 이전과 다른 견해를 제시하고 있다 본 과의 분류 및 계통학적 연구에서, Clausen (1938)은 3개 (Hasebe et al., 1995; Pryer et al., 2001; Smith et al., 2006; 의 속 Botrychium (s.l.), Helminthostachys, Ophioglossum (s.l.) Schuettpelz and Pryer, 2008). 그리고 양치식물군에는 솔잎 을 설정하고 Botrychium속에 subgenus Osmundopteris (Milde) 난강 (Psilotopsida), 속새강 (Equisetopsida), 마라티아강 R.T. Clausen, subgenus Sceptridium (Lyon) R.T. Clausen, (Marattiopsida) 및 고란초강(Polypodiopsida)의 4 강(class)을 subgenus Eubotrychium (Milde) R.T. Clausen등 3개의 아속 두면서, 솔잎난강에 고사리삼과 (Ophioglossaceae)와 솔잎 을 규정한 바 있다(Table 1). 그리고 Ophioglossum속에는 난과 (Psilotaceae)를 두어, 두 과가 계통적으로 가장 가까 subgenus Euophioglossum (Prantl) R.T. Clausen, subgenus 운 것으로 판단한 바 있다. Rhizoglossum (C. Presl) R.T. Clausen, subgenus Cheiroglossa 고사리삼과 식물은 특이한 영양엽편과 포자엽편의 형 (C. Presl) R.T. Clausen, subgenus Ophioderma (Blume) R.T. 태와 위치, 환대가 없는 포자낭 및 권상개엽을 하지 않는 점 Clausen의 4개의 아속 등 전체적으로 1과 3속 7아속을 설 등으로 다른 양치류와 구분되어 뚜렷한 단계통군으로 인식 정하였다(Table 1). 한편 Nishida(1952)는 Clausen의 Ophio- 되고 있으며 (Wagner, 1990; Sun et al., 2001), 한반도의 12 glossaceae (s.l.)를 Ophioglossaceae, Helminthostachyaceae 및 종을 포함하여 전세계적으로 약 80여종이 분포하고 있다 Botrychiaceae로세분하고, Ophiglossaceae (s.s.) 아래에 Ophio- (Mabberley, 1990; Lee, 1996). 과내 속은 전통적으로 고사리 glossum (s.s.), Ophioderma (Blume) Endl., Cheiroglossa C. 삼속 (Botrychium Sw.), 나도고사리삼속 (Ophioglossum L.), Presl, Rhizoglossum C. Presl의 4속을, Helminthostachyaceae Helminthostachys Kaulf. 그리고 제주도에서 기재된 제주고 에 Helminthostachys를, Botrychiaceae에 Botrychium (s.s.), 사리삼속 (Mankyua Sun et al.)의 4개로 구분되며 (Sun et al., 2001), Sceptridium Lyon, Osmundopteris (Milde) Small의 3 속 등 전 이들의 형태적 특징은 다음과 같다 (Clausen, 1938; Eames, 체적으로 3과 8속으로 구분하였다(Table 1). 이어서 Kato 1976; Gilford and Foster, 1989; Kato, 1995; Sun et al., 2001). (1987)는 형태 및 해부학적 특성을 바탕으로 Clausen의 고사리삼속은 영양엽편과 포자엽편이 우상으로 분지 Botrychium (s.l.)을 Botrychium (s.s.), Botrypus Michx, Sceptri- 하고 엽맥은 유리맥이며 지하경은 직립하는 특징을 지닌 dium Lyon 및 Japanobotrychium Masam.의 4개의 속으로 세 다. 반면에 Helminthostachys는 영양엽편이 세 부분으로 나 분하여 본 과를 Ophioglossum (s.l.)과 Helminthostachys를 포 누어지며 각 부분은 다시 우상으로 갈라지고 엽맥은 고사 함하여 6속으로 구분하였다(Table 1). 반면 Wagner (1990) 리삼속과 마찬가지로 유리맥을 이룬다. 그러나 고사리삼 는 Ophioglossaceae (s.l.)를 Botrychiaceae와 Ophioglossaceae 속과는 달리 짧은 자루에 수 개의 포자낭이 달려 전체적 의 두 과로 구분하면서, Botrychiaceae내에 subfamily Botry- 으로 원통 모양의 포자낭수가 생기며 지하경은 횡주하는 chioideae C. Presl과 subfamily Helminthostachyoideae C. Table 1. Comparison of infrafamilial classification system of Ophioglossaceae (s.l.). Clausen (1938) Nishida (1952) Kato (1987) Wagner (1990) Botrychiaceae Botrychiaceae Botrychium (s. l.) Botrychioideae C. Presl Botrychium (s. s.) subg. Eubotrychium (Milde) R.T. Clausen Botrychium (s. s.) Sceptridium Lyon subg. Sceptridium (Milde) R.T. Clausen Botrychium (s. s.) Sceptridium Lyon Osmundopteris (Milde) Small subg. Osmundopteris (Milde) R.T. Clausen Sceptridium Lyon Japanobotrychium Masam. Helminthostachyaceae Helminthostachys Japanobotrychium Masam. Botrypus Michx Helminthostachys Ophioglossum (s. l.) Botrypus Michx Helminthostachyoideae C. Presl Ophioglossaceae (s. s.) subg. Euophioglossum (Prantl) R.T. Clausen Helminthostachys Helminthostachys Ophioglossum (s. s.) subg. Rhizoglossum (C.Presl) R.T. Clausen Ophioglossum (s. l.) Ophioglossaceae Rhizoglossum C. Presl subg. Cheiroglossa (C.Presl) R.T. Clausen Ophioglossum (s. s.) Cheiroglossa C. Presl subg. Ophioderma (Blume) R.T. Clausen Cheiroglossa Ophioderma (Blume) Endl. Ophioderma 한국식물분류학회지 제39권 3호 제주고사리삼을 중심으로한 고사리삼과 식물의 계통 137 Presl의 두 아과를 설정하고, subfamily Botrychioideae에 렬한 후 최종적으로 MacClade 프로그램으로 직접 확인하 Botrychium (s.s.), Scepridium, Botrypus, Japanobotrychium 등 였다. 군외군은 고사리삼과와 유연관계가 깊다고 알려진 4개의 속을, subfamily Helminthostachyoideae에 Helmin- 마라티아과 (Marratiaceae)의 2종 (Angiopteris evecta (G. thostachys속을, 그리고 Ophioglossaceae에 Ophioglossum Forst.) Hottm., Angiopteris parvifolia Ching & Fu)과 Pryer et (s.s.), Ophioderma, Cheiroglossa 등의 3개 속 등 전체적으로 al. (2001)이 고사리삼과와 계통적 유연관계가 밀접한 것 2과 8속을 설정한 바 있다(Table 1). 아울러 Sun et al. (2001)은 으로 판단한 솔잎난과 (Psilotaceae)의 솔잎난 (Psilotum 제주도에서 제주고사리삼속 (Mankyua Sun et al.)을 새롭게 nudum (L.) Griseb.)을 선정하였다. 기재한 바 있다. 최근의 분자계통학적 연구에 있어서 Hasebe PAUP program (Swofford, 2002)의 parsimony analysis를 이 et al.(1993, 1995)은 엽록체 rbcL 유전자 염기서열분석 결 용하여 분지 분석을 수행하였으며, 분석 방법은 Branch- 과를 바탕으로 본 과의 일부 속에 관한 계통을 언급한 바가 and-bound search를 이용하고, 그에 따른 option으로 있었으나 취급한 종 수가 제한적이어서 과 전체의 계통은 ACCTRAN, MULPARS, furthest를 이용하였다. 또한 각 분 논의되지 못하였다. 반면 Hauk et al. (2003)은 Mankyua를 제외 계의 지지 정도를 알아보기 위하여 bootstrap분석 (Felsenstein, 한 Botrychium (s.l.), Ophioglossum (s.l.) 및 Helminthostachys를 포 1985)을 수행하였다. Bootstrap 분석 방법은 1000회를 반복 함하는 과내 36종에 대한 분자계통 및 형태 분석을 바탕 하였으며 이를 통해 각 분계의 통계적 지지도를 추정하였 으로 과내 계통과 기존의 체계에 대하여 논의한 바 있다. 다. 또한 two-parameter method (Kimura, 1980)로 계산된 염 본 연구에서는 현재까지 최근 새롭게 기재된 제주고사 기변이 값을 기초로 한 neighbor-joining tree (NJ)를 산출하 리삼속을 포함한 계통학적 연구가 이루어진 바가 없는 점 였다 (Saitou and Nei, 1987; Farris et al., 1996). 을 고려하여, 제주고사리삼속의 과내에서의 계통학적인 포자의 관찰은 초산분해과정 (Erdtman, 1954, 1966)을 이 위치를 파악하고 나아가 제주고사리삼속을 포함하는 과 용하여 분류군별로 최소 30개의 정상적인 포자를 선택한 전체의 계통을 논의하고자 하였다. 후 극축면의 직경, 적도면의 길이 및 발아구 (laesura)의 길 이 등의 정량적인 형질을 측정하여 각 형질의 최소값, 최 재료 및 방법 대값, 평균 및 표준편차를 구하였다. 재료는 2000년 3월부터 2002년 12월에 걸쳐 국내 및 러 결 과 시아, 일본, 중국 등지에서 채집되었으며, 전북대학교 석 엽표본관 (JNU), 서울대학교 석엽표본관 (SNU), 영남대학 엽록체 rbcL 염기서열의 분석: Botrychium 14종과 Ophio- 교 표본관 (YNUH), 제주도 난대산림연구소 및 중국과학 glossum 12종, Helminthostachys속 1종과 Mankyua속 1종 그 원 북경식물표본관 (PE)에 소장된 표본도 이용하였다. 채 리고 군외군으로 Angiopteris 2종과 솔잎난 (Psilotum 집된 개체는 석엽표본으로 제작하여 전북대학교 생물과 nudum) 1종 등 총 31종에 대한 엽록체 DNA의 rbcL 부위의 학부 석엽표본관 (JNU)에 보관하였으며 일부는 생육하는 염기서열을 결정하였다. 과정을 관찰하기 위해 온실에 이식 재배하였다. 엽록체 전체 rbcL의 길이는 1,314 bp로 이 중 314개가 계통학적 rbcL 유전자 염기서열 분석과 포자 관찰에 사용된 재료의 으로 해상도를 가자는 것으로 나타났다. 또한 Kimura's 확증표본 목록과 아울러 gene bank에서 염기서열이 입수 two parameter model로 계산한 염기 서열의 거리는 0에서 된 종류는 Table 2에 나타난 바와 같다. 0.183으로 나타났으며, 군내군에서는 B. boreale와 O. DNA의 추출은 DNeasy Plant Mini Kit (QIAGEN, pendulum과의 거리가 0.149로 가장 큰 것으로 나타났다. Germany)를 이용하였다. 석엽표본의 잎을 대상으로 한 일 계통수의 길이는 811 단계로, 단독파생형질을 제외한 부 분류군은 Doyle and Doyle (1987)의 CTAB 방법을 이용 Consistency index (CI)가 0.5338이고 Retention index(RI)가 하였고 추출된 DNA는 Ethanol 침전법으로 정제하였다.
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    Acta Box. Neerl. 29 (2/3), May 1980, p. 199-202. Observations on Helminthostachys Kaulfuss (Ophioglossales) 1 2 H.K. GoswamiI and Sharda Khandelwal Department of Botany, Government Science College, Gwalior, M. P. India SUMMARY New observations on the fern genus Helminthostachys Kaulfuss (Ophioglossales) are presented. Tetrarchy ofthe stele in the root is confirmed. The stomata in Helminthostachysare similar tothose in Ophioglossum palmatum.The unusual structure of the vegetative laciniae found at the distal end of In each sporangiophore is again emphasized. the rhizomes the presence of a periderm could be demonstrated. 1. INTRODUCTION Kaulfuss is of the in the Helminthostachys a monotypic genus Ophioglossales eusporangiate ferns. H. zeylanica is found in Ceylon, India, Malay Peninsula, Indonesia, China, Japan, Australia, Philippines, New Caledonia, New Guinea and the Solomon Islands (Beddome 1892, Eames 1936, Panigrahi & Dixit 1969). Numerous workers have made morphological and anatomical studies on the Farmer & Freeman Gwynne- genus (Prantl 1883, Boodle 1899, 1899, Vaughan 1902, Lang 1915, Bower 1926, 1935, Ogura 1938, 1972, Nishida 1956); they have mostly emphasized its similarity to Botrychum Schwarz and Ophioglossum L., despite the significant differences in the fertile spikes of the three genera. is difficult offer of the of It as yet to a morphological explanation rosette vegetative laciniae produced by the sporangiophore, a structure which is unique in the entire plant kingdom (Scott 1923, Bower 1926, 1935) although they are believed to be comparable to growths found in the carboniferous fern Botryo- pteris (Sporne 1970, Bierhorst 1971). this based andanatomical studies the its In paper, on morphological on genus, uniqueness is reemphasized and the suggestion is made that an extensive com- is this parative study needed ofspecimens of monotypic pteridophyte genusfrom different because variables localities, particularly some phenotypic may be sug- gestive of genetic and/or adaptive changes.
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  • Conservation Status of the Endemic Fern Mankyua Chejuense (Ophioglossaceae) on Cheju Island, Republic of Korea
    Oryx Vol 38 No 2 April 2004 Short Communication Conservation status of the endemic fern Mankyua chejuense (Ophioglossaceae) on Cheju Island, Republic of Korea Chul Hwan Kim Abstract Mankyua chejuense, a fern endemic to Cheju Endangered on the IUCN Red List. For conservation Island, Republic of Korea, which lies 120 km south of the of the species it needs to be included on the national Korean Peninsula, appears to be restricted to five extant threatened species list, and its habitat designated as subpopulations in the north-east of the Island, with a an ecological reserve. Intensive surveys are required total population of c. 1,300 individuals. Major threats to in order to establish whether there are any other extant the existence of the species include shifting cultivation, subpopulations of the species, and the presently known plantation, overuse of basaltic rocks that are part of the subpopulations require long-term monitoring and continuous protection. species’ microhabitat, farming and pasturage, and the construction of roads and golf courses in lowland areas. Keywords Cheju Island, Critically Endangered, The information currently available for the species endemic, fern, Mankyua chejuense, Ophioglossaceae, indicates that it should be categorized as Critically Republic of Korea. The 1,824 km2 Cheju Island is located 140 km south of the number of individuals. In this paper I report an approxi- Korean Peninsula. It is of volcanic origin and the highest mate total count of M. chejuense, examine its conservation peak, Halla Mountain, rises to an altitude of 1,950 m. The status, identify factors threatening the species’ survival, island has three main landscape types: lowland areas and propose an IUCN Red List status for the species.
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  • (OUV) of the Wet Tropics of Queensland World Heritage Area
    Handout 2 Natural Heritage Criteria and the Attributes of Outstanding Universal Value (OUV) of the Wet Tropics of Queensland World Heritage Area The notes that follow were derived by deconstructing the original 1988 nomination document to identify the specific themes and attributes which have been recognised as contributing to the Outstanding Universal Value of the Wet Tropics. The notes also provide brief statements of justification for the specific examples provided in the nomination documentation. Steve Goosem, December 2012 Natural Heritage Criteria: (1) Outstanding examples representing the major stages in the earth’s evolutionary history Values: refers to the surviving taxa that are representative of eight ‘stages’ in the evolutionary history of the earth. Relict species and lineages are the elements of this World Heritage value. Attribute of OUV (a) The Age of the Pteridophytes Significance One of the most significant evolutionary events on this planet was the adaptation in the Palaeozoic Era of plants to life on the land. The earliest known (plant) forms were from the Silurian Period more than 400 million years ago. These were spore-producing plants which reached their greatest development 100 million years later during the Carboniferous Period. This stage of the earth’s evolutionary history, involving the proliferation of club mosses (lycopods) and ferns is commonly described as the Age of the Pteridophytes. The range of primitive relict genera representative of the major and most ancient evolutionary groups of pteridophytes occurring in the Wet Tropics is equalled only in the more extensive New Guinea rainforests that were once continuous with those of the listed area.
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  • Morphological Variations of Fertile Spike in Helminthostachys Zeylanica (L.) Hook
    HACQUETIA 11/2 • 2012, 271–275 DOi: 10.2478/v10028-012-0007-0 MorphologIcAl vAriatIons of fertIle spike In HelmintHostacHys zeylanica (l.) hooK Tapas K. CHaKRabORTy1,2,*, Sandip Dev CHauDHuRi2 & Jayanta CHOuDHuRy2 Abstract The evolutionary history of Ophioglossaceae is enigmatic mainly because fossils of the family trace back only from the earliest Tertiary. Phylogenetic analyses indicate that Helminthostachys is sister to the broadly defined Botrychium. Generally the sporophore of Botrychium is a pinnately compound, whereas it is simple in Hel- minthostachys. Here examples of different Helminthoistachys are represented which show double or triple spikes with some variations. Plants showing variations in their spike morphology are also grown normally. Variations of Helminthostachys spike morphology indicate a tendency to form a compound sporophore structure and in that way have a strong relationship with Botrychium. Key words: Helminthostachys zeylanica, pteridophyte, Ophioglossaceae, fertile spike. Izvleček: Evolucijski razvoj družine kačjejezikovk (Ophioglossaceae) je precej skrivnosten, saj prve fosilne ostanke pred- stavnikov družine najdemo šele iz zgodnjega terciarja. Filogenetske raziskave kažejo, da je monotipični rod Helminthostachys sestrski sicer širše definiranemu rodu mladomesečin (Botrychium). Na splošno je fertilni ali plodni del sporotrofofila pri rodu Botrychium pernato deljen, medtem ko je pri rodu Helminthostachys valjast in enostaven. V študiji so predstavljeni različni osebki vrste Helminthostachys zeylanica z enim ali več plodnimi izrastki. Osebki, ki kažejo na variacijo v obliki trosnih izrastkov rastejo v naravnem okolju. Različna morfologija plodnega-trosnega dela sporotrofofila pri rodu Helminthostachys nakazuje tendenco k tvorbi pernato deljenih trosnih izrastkov, kar je znak sorodnosti omenjenega rodu z rodom Botrychium. Ključne besede: Helminthostachys zeylanica, praprotnice, Ophioglossaceae, plodni izrastek.
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  • Fern Classification
    16 Fern classification ALAN R. SMITH, KATHLEEN M. PRYER, ERIC SCHUETTPELZ, PETRA KORALL, HARALD SCHNEIDER, AND PAUL G. WOLF 16.1 Introduction and historical summary / Over the past 70 years, many fern classifications, nearly all based on morphology, most explicitly or implicitly phylogenetic, have been proposed. The most complete and commonly used classifications, some intended primar• ily as herbarium (filing) schemes, are summarized in Table 16.1, and include: Christensen (1938), Copeland (1947), Holttum (1947, 1949), Nayar (1970), Bierhorst (1971), Crabbe et al. (1975), Pichi Sermolli (1977), Ching (1978), Tryon and Tryon (1982), Kramer (in Kubitzki, 1990), Hennipman (1996), and Stevenson and Loconte (1996). Other classifications or trees implying relationships, some with a regional focus, include Bower (1926), Ching (1940), Dickason (1946), Wagner (1969), Tagawa and Iwatsuki (1972), Holttum (1973), and Mickel (1974). Tryon (1952) and Pichi Sermolli (1973) reviewed and reproduced many of these and still earlier classifica• tions, and Pichi Sermolli (1970, 1981, 1982, 1986) also summarized information on family names of ferns. Smith (1996) provided a summary and discussion of recent classifications. With the advent of cladistic methods and molecular sequencing techniques, there has been an increased interest in classifications reflecting evolutionary relationships. Phylogenetic studies robustly support a basal dichotomy within vascular plants, separating the lycophytes (less than 1 % of extant vascular plants) from the euphyllophytes (Figure 16.l; Raubeson and Jansen, 1992, Kenrick and Crane, 1997; Pryer et al., 2001a, 2004a, 2004b; Qiu et al., 2006). Living euphyl• lophytes, in turn, comprise two major clades: spermatophytes (seed plants), which are in excess of 260 000 species (Thorne, 2002; Scotland and Wortley, Biology and Evolution of Ferns and Lycopliytes, ed.
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  • Biology of Ophioglossum L
    Bionature, 27 (1 & 2), 2007 : 1-73 © Bionature BIOLOGY OF OPHIOGLOSSUM L. H. K. GOSWAMI ABSTRACT Ophioglossales are the natural group of early vascular plants which exhibit the most simple and most complicated combinations of characters comparable to bryophytes, pteridophytes, progymnosperms, gymnosperms and angiosperms. Essentially, pteridophytes these plants are often referred and classified as ferns. However, there are some fundamental differences which should not justify their present alliance. The chief "genetic loss" in plants of this group can be presumed to be the loss of capability of producing sclerenchyma. Also, the sporangia are unlike ferns; they do not have an annulus and are supplied with vascular tissue. Additionally, absence of circinate vernation and presence of periderm (in about 22% of Ophioglossum population) make them "unlike ferns". The conventionally recognised three genera, Botrychium, Helminthostachys and Ophioglossum constitute a single family Ophioglossaceae of the order Ophioglossales. Nevertheless, intergeneric differences are so pronounced that recognition of three separate families viz. Botrychiaceae, Helminthostachyaceae and Ophioglossaceae by some taxonomists are quite justified. Botrychium and Ophioglossum are further divided to have subgenera; Botrychium has Sceptridium, Eubotrychium and Osmundopteris, while Ophioglossum has two, viz. Ophioglossum and Ophioderma. Population cytogenetic studies have been carried out chiefly from the localities where more than one species of Ophiglossum grow. Repeated meiotic studies have also been carried out from populations of single or isolated species of Ophioglossum and monotypic Helminthostachys. Numerous teratologies of genetic importance have been described. Role of natural selection is being assessed. Lately, a new specis O. eliminatum is being suspected to have been arisen by natural hybridization and chromosomal elimination.
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  • A Review on the Potential Uses of Ferns M
    Ethnobotanical Leaflets 12: 281-285. 2008. A Review on the Potential Uses of Ferns M. Mannar Mannan, M. Maridass* and B.Victor Animal Health Research Unit, St. Xavier’s College (Autonomous) Palayamkottai, Tamil Nadu – 627002 *Corresponding Author: Dr. M. Maridass, DST-SERC-Young Scientist Animal Health Research Unit St. Xavier’s College (Autonomous), Palayamkottai, Tamil Nadu – 627002. Email: [email protected] Issued 24 May 2008 Introduction Man has been using plants as a source of food, medicines and many other necessities of life since ancient times. Even to this day the primitive tribal societies that exist depend on the plant life in their surroundings. Though there were investigations of the edible economic values of the higher plants, especially the pteridophytes and angiosperms have been unfortunately ignored. The pteridophytes are used in Homoeopathic, Ayurvedic, Tribal and Unani medicines and provides food, insecticides and ornamentations. Ferns used as food With very few exception ferns have not been widely used as a source of food. The fern stems, rhizomes, leaves, young fronds and shoots and some whole plants are used for food. Tree ferns have often been used as food and starch in Hawaii. Also, ferns are supposed to increase milk production when fed to cows in Sicily. The young fronds and underground stem of the fern Asplenium ensiforme are used for food by hilly tribes. In Malaysia, Blechnum orientalis L., rhizome is eaten and whole plant is used as feed and as poultice in boil. The fronds of Ceratopteris thalictroides are used as a vegetable. The young fronds of Diplazium esculentum are eaten either as salad or as vegetable after cooking.
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  • OPHIOGLOSSACEAE 1. BOTRYCHIUM Swartz, J. Bot
    This PDF version does not have an ISBN or ISSN and is not therefore effectively published (Melbourne Code, Art. 29.1). The printed version, however, was effectively published on 6 June 2013. Zhang, X. C., Q. R. Liu & N. Sahashi. 2013. Ophioglossaceae. Pp. 73–80 in Z. Y. Wu, P. H. Raven & D. Y. Hong, eds., Flora of China, Vol. 2–3 (Pteridophytes). Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. OPHIOGLOSSACEAE 瓶尔小草科 ping er xiao cao ke Zhang Xianchun (张宪春)1, Liu Quanru (刘全儒)2; Norio Sahashi3 Plants perennial, mostly terrestrial, rarely epiphytic, usually small and fleshy, lacking sclerenchyma. Roots lacking root hairs, unbranched or with a few narrow lateral branches [rarely dichotomously branched], fibrous or fleshy, sometimes producing vegetative buds. Rhizome mostly erect, less often horizontal, rarely branched, eustelic, glabrous or hairy. Fronds 1 to few per plant, monomorphic, vernation nodding (not circinate), erect or folded, stipe base dilated, clasping, forming open or fused sheath surrounding successive leaf buds; buds glabrous or with long, uniseriate hairs; common stipe usually dividing into sterile, laminate, photosynthetic portion (trophophore) and fertile, spore-bearing portion (sporophore); sterile lamina ternately or pinnately compound to simple, rarely absent, glabrous or with scattered, long, uniseriate hairs, especially on stipe and rachis; veins anastomosing or free, pinnate, or palmate. Sporophores 1 per frond [rarely more], spikelike or pinnately branched; sporangia exposed or embedded, some- times clustered on very short lateral branches, wall 2 cells thick, annulus absent; spores many (> 1000) per sporangium, globose- tetrahedral, trilete, thick-walled, surface rugate, tuberculate, baculate (with projecting rods usually higher than wide), sometimes joined in delicate network, mostly with ± warty surface.
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  • Taxonomic Significance of Morphological Characters of Spores
    Review of Palaeobotany and Palynology 252 (2018) 77–85 Contents lists available at ScienceDirect Review of Palaeobotany and Palynology journal homepage: www.elsevier.com/locate/revpalbo Taxonomic significance of morphological characters of spores in the family Ophioglossaceae (Psilotopsida) Natalia Olejnik a,⁎, Zbigniew Celka a,PiotrSzkudlarza, Myroslav V. Shevera b a Department of Plant Taxonomy, Faculty of Biology, Adam Mickiewicz University in Poznań, Umultowska 89, 61-614 Poznań,Poland b Department of Systematics and Floristic of Vascular Plants, M. G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, Kyiv, Ukraine article info abstract Article history: Primary and secondary ornamentation of spores of ferns of the family Ophioglossaceae are important characters, Received 16 September 2017 used in the taxonomy of this group. Considering the small number of published data on those characters in the Received in revised form 7 February 2018 Ophioglossaceae from Central and Eastern Europe, this study aimed (1) to describe morphological characters Accepted 19 February 2018 of spores of Botrychium and Ophioglossum species and to assess their taxonomic significance; (2) to analyse var- Available online 21 February 2018 iation in spore size between and within species of these genera, based on specimens from various habitats and fi Keywords: geographic locations; and (3) to create a key to species identi cation based on the diagnostic characters of the Class Psilotopsida spore ornamentation. We examined spores of 6 species from 16 localities in Central and Eastern Europe. Results Key to species identification of cluster analysis based on morphological characters of spores indicate that the species form well-defined Morphology groups, partly reflecting the systematics of the Ophioglossaceae.
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  • The First Asian Plant Conservation Report
    The Convention on Biological Diversity The First Asian Plant Conservation Report A Review of Progress in Implementing the Global Strategy for Plant Conservation (GSPC) Published by Chinese National Committee for DIVERSITAS (CNC- DIVERSITAS), Beijing, China Copyright: © 2010 Chinese National Committee for DIVERSITAS Resources: Reproduction of this publication for educational or other non- commercial purposes is authorized without prior written permission from the copyright holder provided the source is fully acknowledged. Reproduction of this publication for release or other commercial purposes is prohibited without prior written permission from the copyright holder. This publication has been made possible by funding from CNC-DIVERSITAS Layout by: Bing Liu and Yinan Liu Produced by: Beijing Changhao Printing Co., Ltd. Citation: Keping Ma et al. (2010). The First Asian Plant Conservation Report. Beijing, China. 66pp. Available from: Secretariat of Chinese National Committee for DIVERSITAS Address: No.20, Nanxincun, Xiangshan, Beijing 100093, China Tel: 86-10-62836603 Fax: 86-10-82591781 E-mail: [email protected] Website: http://www.cncdiversitas.org/ Contents Forward by Dr. Peter H. Raven ………………………………………………………2 Forward by Ms. Aban Marker Kabraji …………………………………………………3 Preface …………………………………………………………………………………4 Executive Summary ……………………………………………………………………6 Section 1: Brief introduction of GSPC …………………………………………………9 Section 2: Overview of Asia …………………………………………………………10 Section 3: Key features of plant diversity in Asia……………………………………11 Section
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  • Studies on the Cytology and Phylogeny of the Pteridophytes VI. Observations on the Ophioglossaceae
    1958 291 Studies on the Cytology and Phylogeny of the Pteridophytes VI. Observations on the Ophioglossaceae C. A. Ninan Department of Botany, University College , Trivandrum, India Received January 24, 1958 Introduction The Ophioglossaceae is a "very distinctive and circumscribed family" of primitive megaphyllous Pteridophytes consisting of three living genera, Ophioglossum, Botrychium and Helminthostachys. Without any known fossil record and with very distinctive features, the three genera constitute a natural family, their common character being the possession of the fertile spike. This family enjoys world-wide distribution and consists of one hundred species (Carl Christensen 1905-1934), the monotypic Helmin thostachys being restricted to the Australian and Indo-Malayan regions. Bower (1926) recognizes 78 species in this family while Clausen (1938) reduces them to 50. The three genera are typically of the eusporangiate type and combine several points of interest in cytology, phylogeny and evolution. The genus Ophioglossum is typical of the family and is perhaps the most ancient of all living ferns. It consists of 56 species according to Christensen's index (Bower and Clausen recognize only 43 and 26 species respectively). Most of them are ground growing forms while two species, O. pendulum and O. palmatum are epiphytic. Over a dozen species are indigenous to India and are found distributed in a variety of habitats. Botrychium is represented by 43 species (Christensen 1905-1934). Bower and Clausen recognize 34 and 23 species respectively for this genus. In the tropics this genus is usually confined to higher elevations. The only species of the genus Helminthostachys (H. zeylanica) is usually found to occur in low lands or river sides which get inundated.
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  • A Classification for Extant Ferns
    55 (3) • August 2006: 705–731 Smith & al. • Fern classification TAXONOMY A classification for extant ferns Alan R. Smith1, Kathleen M. Pryer2, Eric Schuettpelz2, Petra Korall2,3, Harald Schneider4 & Paul G. Wolf5 1 University Herbarium, 1001 Valley Life Sciences Building #2465, University of California, Berkeley, California 94720-2465, U.S.A. [email protected] (author for correspondence). 2 Department of Biology, Duke University, Durham, North Carolina 27708-0338, U.S.A. 3 Department of Phanerogamic Botany, Swedish Museum of Natural History, Box 50007, SE-104 05 Stock- holm, Sweden. 4 Albrecht-von-Haller-Institut für Pflanzenwissenschaften, Abteilung Systematische Botanik, Georg-August- Universität, Untere Karspüle 2, 37073 Göttingen, Germany. 5 Department of Biology, Utah State University, Logan, Utah 84322-5305, U.S.A. We present a revised classification for extant ferns, with emphasis on ordinal and familial ranks, and a synop- sis of included genera. Our classification reflects recently published phylogenetic hypotheses based on both morphological and molecular data. Within our new classification, we recognize four monophyletic classes, 11 monophyletic orders, and 37 families, 32 of which are strongly supported as monophyletic. One new family, Cibotiaceae Korall, is described. The phylogenetic affinities of a few genera in the order Polypodiales are unclear and their familial placements are therefore tentative. Alphabetical lists of accepted genera (including common synonyms), families, orders, and taxa of higher rank are provided. KEYWORDS: classification, Cibotiaceae, ferns, monilophytes, monophyletic. INTRODUCTION Euphyllophytes Recent phylogenetic studies have revealed a basal dichotomy within vascular plants, separating the lyco- Lycophytes Spermatophytes Monilophytes phytes (less than 1% of extant vascular plants) from the euphyllophytes (Fig.
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