杉科的染色体资料及在系统演化研究中的作用

杉科共由10属(包括金松属)、20种(变种)组成。本文整理了19种(占95％,隶10属)植物的染色体数目和16种(占80％,隶9属)的核型资料,核型的模式图如图1所示。通过对这些细胞学资料的分析,笔者支持2n=20、x=10的金松属从杉科(2n=22、x=11)分立成金松科。根据其他各属间的亲缘关系,本作者认为可以把它们分隶于5个亚科:Ⅰ.柳杉亚科:Cryptomerioideae(Cryptomeria);Ⅱ.落羽杉亚科Taxodioideae(Glyptostroous,Taxodium);Ⅲ.红杉亚科Sequoideae(Metasequoia,Sequoiadendron,Sequoia);Ⅳ.杉木亚科Cunninghamioideae(Cunninghamia,还可能有:Athrotaxis);Ⅴ.台湾杉亚科Taiwainoideae(Taiwania)。这些亚科和属的进化水平依序渐增,它们分别位于进化路线A(亚科Ⅰ、Ⅱ、Ⅴ)和进化路线L(亚科Ⅲ、Ⅳ)上。这些结果是前人的演化系统所没有涉及的,表明了染色体资料在杉科的系统演化研究中起着重要的作用。     

杉科的细胞分类学和系统演化研究

根据杉科的核型资料,本文(1)提出“1B”可能是一个新的高等植物核型类型;(2)讨论了各属的有关分类学问题及相互亲缘关系,它们的进化顺序可能是柳杉属、水松属、落羽杉属、水杉属、巨杉属、红杉属、杉木属(密叶杉属与之近缘)、台湾杉属;(3)支持金松属分立成金松科,它可能比杉科各属原始;(4)红杉(AAAABB)的亲本可能是二个古代种水杉”和“巨杉”,它们的直接后裔或留下的近缘是水杉和巨杉;(5)杉科存在A和L两条进化路线,前者包括柳杉属、水松属、落羽杉属、台湾杉属,后者包括水杉属、巨杉属、红杉属、杉木属(密叶杉属);(6)提出一个杉科新系统(包括一个新亚科)：I．柳杉亚科(柳杉属),II．落羽杉亚科(水松属、落羽杉属),III．红杉亚科(冰杉属、巨杉属、红杉属),Ⅳ.杉木亚科(杉木属、密叶杉属),V．台湾杉亚科,新亚科(台湾杉属)。本文还对前人的杉科系统作了讨论。     

杉科的细胞分类学和系统演化研究

根据杉科的核型资料,本文(1)提出“1B”可能是一个新的高等植物核型类型;(2)讨论了各属的有关分类学问题及相互亲缘关系,它们的进化顺序可能是柳杉属、水松属、落羽杉属、水杉属、巨杉属、红杉属、杉木属(密叶杉属与之近缘)、台湾杉属;(3)支持金松属分立成金松科,它可能比杉科各属原始; (4)红杉(AAAABB)的亲本可能是二个古代种“水杉”和“巨杉”,它们的直接后裔或留下的近缘是水杉和巨杉;(5)杉科存在A和L两条进化路线,前者包括柳杉属、水松属、落羽杉属、台湾杉属;后者包括水杉属、巨杉属、红杉属、杉木属(密叶杉属);(6)提出一个杉科新系统(包括一个新亚科):Ⅰ.柳杉亚科(柳杉属),Ⅱ.落羽杉亚科(水松属、落羽杉属),Ⅲ.红杉亚科(水杉属、巨杉属、红杉属),Ⅳ.杉木亚科(杉木属、密叶杉属),Ⅴ.台湾杉亚科,新亚科(台湾杉属)。本文还对前人的杉科系统作了讨论。     

杉科、柏科的系统发生关系研究进展

杉科、柏科是松柏类裸子植物中的重要类群,其系统分类研究一直是裸子植物的研究热点之一.但是杉科与柏科之间及其内部各属之间的系统发生关系却一直存在争议.一般认为杉科、柏科单独成科.近年来的分子系统学及分支系统学研究结果证实:除了金松属以外,杉科和柏科为一单系群,应合并为一个科Cupres-saceae sensu　lato（广义柏科）,其主要分支类群的系统发生关系也已经基本确立,而金松属则单独成立金松科.     

秃杉的细胞学研究

本文首次报道秃杉的核型为K(2n)=22=16m+6sm, 3号染色体具“长着丝点区 域”。似具比同属的台湾杉(18m+4sm)进化的趋势。它们均为"2B"。从国产杉科植物的 核型类型看,似见"2B"的台湾杉属与"1B"的杉木属和"1A"的水杉属、水松属、柳杉属的亲缘关系依序由近到远并比它们进化。本作者还观察了秃杉的核仁数目和多微核现象。     

辽宁早白垩世化石木一新属——原始金松型木（Protosciadopityoxylon）

报道了辽宁早白垩世化石木的一个新属—原始金松型木 (Protosciadopityoxylongen .nov .)。该属化石以辽宁原始金松型木 (Protosciadopityoxylonliaoningensegen .etsp .nov .)为代表。文中对新属新种进行了描述 ;讨论了新属与现存金松属 (Sciadopitys)及化石金松型木属 (Sciadopityoxylon)的关系 ;并与其相似的原始叶枝杉型木属 (Protophyl locladoxylon)、原始圆孔木属 (Protocircoporoxylon)及异木属 (Xenoxylon)作了比较。     

杉科植物的系统发育分析

本文以形态学为依据,参考其他学科的研究成果,用分支分类方法并结合表征分类方法探讨了杉科植物的系统演化关系,提出了新的分类系统。在分支分类中,金松科被选作外类群。主要根据外类群比较原则、化石原则和一般的演化规律,确定了性状的祖征和衍征,采用最大同步法、综合分析法、演化极端结合法及最小平行进化法共四种方法进行分支分析,选择最简约的分支图作为本文讨论基础。在表征分类中,选取59个性状,利用距离系数和类平均法,对金松属和杉科各属进行了聚类运算,得出表征图。综合两种分析结果,主要结论如下:(1)属间关系:柳杉属是现存杉科植物中最原始的类群。水松属和落羽杉属关系密切,二者与柳杉属近缘。巨杉属和北美红杉属关系密切,是中级进化水平的类群。水杉属与巨杉属和北美红杉属的亲缘关系相对较近。杉木属、密叶杉属和台湾杉属关系密切,是杉科植物中的高级进化类群,其中又以台湾杉属演化水平最高。(2)系统排列:支持金松科的成立,将杉科分成5族,即柳杉族(仅含柳杉属)、落羽杉族(含水松属、落羽杉属)、北美红杉族(含巨杉属、北美红杉属)、水杉族(仅含水杉属)和杉木族(含杉木属、密叶杉属及台湾杉属)。     

生境的破坏或改变对杉科种属分布的影响

杉科共有10个属16种,其中5个是单种属(水杉属、水松属、巨杉属、北美红杉属、金松属),5个是寡种属(杉木属、台湾杉属、柳杉属、落羽杉属、密叶杉属),两者各占50%。分布于北温带地区,多在东亚和北美洲,仅密叶杉属分布于南半球澳大利亚的塔斯马尼亚。植物与生境是一个互相依存的统一体,而生境的破坏或改变对杉科植物种属分布有严重的影响。在美国加利福尼亚洲现今仅有巨杉和北美红杉两个单种属,这是由于火山爆发留下来,现在仅分布于美国加州部分地区。     

密叶杉属的核型分析及其系统位置的探讨

本文首次对特产澳大利亚塔斯马尼亚岛的Athrotaxis cupressoides和Athrotaxis selaginoides进行了核型分析,核型公式分别为2n=22=22m(2SAT)和2n=22=20(2SAT)+2sm,均属Stebbins的1B类型,它们的染色体相对长度组成为22=2L+10M₂+8M_l+2S和22=2L十10M₂+6M_l+4S,后者比前者较为进化。根据密叶杉属和杉科其他各属核型资料的比较分析,它们由原始到进步的顺序可能为：柳杉属、水松属、落羽杉属、水杉属、巨杉属、红杉属,密叶杉属、杉木属和台湾杉属。 密叶属与红杉属、巨杉属和杉木属较为近缘。 这些在以染色体长度比和平均臂比为纵、横坐标的图上得到清楚反映。 根据核型资料,密叶杉属以隶于单型的亚科Arthrotaxoideae较为合适,这也得到形态学、胚胎学、孢粉学和地理学资料的支持。 本文还对前人系统中的密叶杉属位置进行了讨论。     

天南星科花叶万年青属两种植物的核型

报道了天南星科(Araceae)花叶万年青属(Dieffenbachia)两种植物染色体数目及核型,革叶万年青(D.daguensis)品种绿帝皇的核型公式为K(2n)=2x=34=12m 18sm 4st,核型类型为"3B",核型不对称系数As.K%=66.52%。花叶万年青(D.maculata)品种粉黛叶的核型公式为K(2n)=2x=34=2M 28m 4sm,核型类型为"1B",核型不对称系数为AS.K%=55.38%。与同科的较进化的犁头尖属和斑龙芋属比较,花叶万年青属植物的核型较原始些,且天南星科植物的核型极其多样化。     

The Karyotype Analysis of Arthrotaxis (Taxodiaceae) and Its Systematic Position

The present paper deals for the first time with an analysis of the karyotypes of Athrotaxis cupressoides Don and A. selaginoides Don endemic to Tasmania (Australia). Their morphology of somatic chromosomes in seed root-tip cells, chromosome measurements, and diagrams are shown in Plate 1, Table 1 and Fig. 1 respectively, The karyotypic formulas of the two species are 2n = 22 = 22m (2SAT) and 2n = 22 = 20m(2SAT ) + 2sm according to of terminology Lexvan et al (1964). They all belong to IB type of Stebbins’(1971)karyotypic asymmetry which was reported for the first time in the higher plants by Li(1987b). Their chromosome complements are 22 = 2L + 10M₂+ 8M₁+ 2S and 22 =2L+ 10M₂+ 6M₁+ 4S respectively according to the standard defined by Kuo et al. (1972) based on relative length. The karyotype of A. selaginoides is more advanced than that of A. cupressoides. In the light of karyotypic data, the sequence of the taxo-diaceous genera (excl. Sciadopitys) from primitive to advanced may be in the following order: Cryptomeria, Glyptostrobus, Taxodium, Metasequoia, Sequoiadendron, Sequoia, Athrotaxis, Cunninghamia and Taiwania. The genus Athrotaxis is closely related to Sequoia (Sequoiadendron) and Cunninghamia The peculiarity of the karyotype of Athrotaxis deserves the establishment of a new status Arthrotaxoideae (Wettstein) L. C. Li This suggestion is also supported by the data from morphology, embryology, palynology and geography. The family Taxodiaceae is divided into six subfamilies and nine genera, as shown in the following table:————————————————————————————————————————————————— 1. Cryptomerioideae Hida Cryptomeria D. Don 4. Arthrotaxoideae (Wettstein) L. C. Li 2. Taxodioideae Pilger Glyptostrobus Endl. Taxodim Richard Athrotaxis D. Don 3. Sequoideae Saxton metasequoia Miki ex Hu et 5. Cunninghamioideae Hida Cunninghamia Cheng Sequoiadebron Buch. Sequoia Endl. R.Brown 6. Taiwanioideae (Hayata)L. C. Li Taiwanta Hayata————————————————————————————————————————————————— The systematic positions of Athrotaxis in the systems of other authors are dis-cussed too.     

Phylogenetic Analysis of the Family Taxodiaceae

In the present paper,both cladistic analysis and phenetic analysis were conducted to evaluate the phylogenetic relationships of the Taxodiaceae based on an extensive literature review and study of herbarium． In the cladistic analysis,the Sciadopityaceae was chosen as outgroup．The polarity of characters was determined mainly according to outgroup comparison,fossil evidence and generally accepted viewpoints of morphological evolution．By the result of compatibility analysis,character 2(leaf type),which possessed a much higher coefficient than others whether or not its polarity was altered,was deleted． Finally,a data matrix consisting of all the extant nine genera and 24 characters was analyzed using Maximal Same Step Method,Synthetic Method,Evolutionary Extremal Aggregation Method and Minimal Parallel Evolutionary Method,and four cladograms were generated,of which only the most parsimonious one (Fig．1)was presented for discussion. The cladogram shows that the Taxodiaceae are assorted along five lines of evolution： 1)Metasequoia;2)Sequoiadendron,Sequoia;3)Cryptomeria;4)Glyptostrobus and Taxodium;5)Cunninghamia,Athrotaxis and Taiwania． Ten genera(including Sciadopitys)and 59 characters were used in the phenetic analysis．The phenogram(Fig．2)indicates that Sciadopitys is a very distinct group with remote affinity to the other genera,and the Taxodiaceae are divided into four groups：1)Sequoia,Sequoiadendron;2)Athrotaxis,Cunninghamia and Taiwania;3)Cryptomeria,Glyptostrobus and Taxodium;4)Metasequoia． Based primarily on the result of cladistics,with reference to that of phenetics,the main conclusions were drawn as follows：(1)Generic relationships：Cryptomeria should be considered the most primitive genus in the extant groups of the Taxodiaceae. Glyptostrobus and Taxodium, close to Cryptomeria, are sister taxa and relatively primitive groups. Sequoiadendron and Sequoia are closely related and intermediate advanced. Metasequoia is a more or less isolated taxon, relatively close to Sequoiadendron and Sequoia. Cunninghamia. Athrotaxis and Taiwania might represent a single lineage and form a very advanced group, of which Taiwania may be the most specialized. (2) Systematic treatments: The authors support the viewpoint that Sciadopitys should be treated as an independent family, and suggest that the Taxodiaeae should be divided into five tribes. Systematic arrangements are as follows: Taxodiaceae Warming Trib. 1. Cryptomerieae Vierhapper Gen. 5. Sequoia Endl. Gen. 1. Cryptomeria D. Don Trib. 4. Metasequoieae Pilger et Melchior Trib. 2. Taxodieae Benth. et Hook. Gen. 6. Metasequoia Miki ex Hu et Cheng Gen. 2. Glyptostrobus Endl. Trib. 5. Cunninghamieae Zucc. Gen. 3. Taxodium Rich. Gen. 7. Cunninghamia R. Br. Trib. 3. Sequoieae Wettstein Gen. 8. Athrotaxis D. Don Gen. 4. Sequoiadendron Buchholz Gen. 9. Taiwania Hayata     

云杉核型的研究兼论云杉属的进化地位

本文分析了我国特产树种云杉Picea asperata的核型,K(2n)=24=20m+4sm,属2A类型,染色体相对长度组成为2n=24=2L+12M_2+SM_1+2S。云杉属植物(22种、变种)的核型全由臂比小于2的中部和近中着丝粒染色体构成,是较为原始的核型。根据松科各属核型的比较,作者讨论了云杉属的亲缘关系和进化地位,并得到形态学、解剖学、孢粉学、植化学、生化学及古植物学等的支持。     

川滇冷杉的核型分析兼论冷杉属的进化地位

本文分析了我国特产树种川滇冷杉Abies forrestii的核型,K(2n)=24=14 m+8 sin+2 st,属2B类型,染色体相对长度组成为2n=24=2 L+10 M_2+10 M_1+2 S。冷杉属植物(除川滇冷杉)的核型全由中部和近中着丝粒染色体组成,属2A类型(表2)。根据松科各属核型的比较,作者讨论了冷杉属的亲缘关系和进化地位,并得到形态学、解剖学、生化学、古植物学等的支持。     

澳大利亚三种Callitris植物的核型及其系统学意义

对澳大利亚特产的Ｃａｌｌｉｔｒｉｓ属植物Ｃ．ｐｒｅｉｓｓｉｉ,Ｃ．ｖｅｒｒｕｃｏｓａ,Ｃ．ｅｎｄｌｉｃｈｅｒｉ（柏科）的核型进行了分析,后２种的为首次报道。它们的核型公式分别为Ｋ（２ｎ）＝２２＝２２ｍ（２ＳＡＴ）,２２ｍ（２ＳＡＴ）和２２ｍ（６ＳＡＴ）,均属１Ａ核型类型。染色体相对长度组成同是２ｎ＝２２＝１０Ｍ＿２＋１２Ｍ＿１该３种及其他８种Ｃａｌｌｔｒｉｓ属植物一致的核型Ｋ（２ｎ）＝２２ｍ和１Ａ类型的通常被认为是最对称和原始的。因此该属在柏科的系统发育上也许处于相当原始的地位。     

柏科分类和分布：亚科,族和属

柏科Cupressaceae和杉科Taxodiaceae有许多相似之处,近年来不少分类学家主张把两科合并成广义的柏科。原杉科中的金松属Sciadopitys与两科其他属的差异较大,被提升为单种科Sciadopity-aceae。本文根据球果可育种鳞的位置把柏科(狭义)分为2亚科,即上部种鳞不可育的柏木亚科Cupres-soideae和上部种鳞可育的澳洲柏亚科Callitroideae。综合其他形态学和解剖学证据,柏木亚科又分4族,即柏木族Cupresseae(包括:柏术属Cupressus、杂交柏属×Cupressocyparis、扁柏属Chamaecyparis和福建柏属Fokeinia)、侧柏族Thujopsideae(包括:崖柏属Thuja、罗汉柏属Thujopsis和侧柏属Platycladus)、圆柏族Junipereae(包括:圆柏属Juniperus和海参威柏属Microbiota)以及香漆柏族Tetraclineae(包括:翠柏属Calocedrus和香漆柏属Tetraclinis)。澳洲柏亚科又分3族,即澳洲柏族Actinostrobeae(包括:西澳柏属Actinostuobus、澳洲柏属Callitris、智利柏属Fitzroya和杉叶柏属Neocallitropsis)、南非柏族Widdring-toneae(包括:白智利柏属Pilgerodendron、塔斯曼柏属Diselma和南非柏属Widdringtonia)以及甜柏族Libocedreae(包括:甜柏属Libocedrus、巴布亚柏属Papuacedrus和南美柏属Austrocedrus)。柏科21个属的地理分布可划分为5种类型,即:(     

水松的细胞学研究

本文报道了水松的核型公式K(2n)=22=22m,为“1A”类型。染色体相对长度组成为2n=22=2L+4M_2+16M_1。8号染色体具长着丝点区域,这是核型的一个特征。与近缘的国产种柳杉和水杉相比较。三者由原始到进化的顺序可能为(柳杉、水松)、水杉,水松与柳杉最接近,水杉和水松较近缘。本文还计算了水松的染色体体积。     

Taxonomy of the Cupressaceae: Subfamilies,Tribes and Genera

Cupressaceae and Taxodiaceae have recently been merged under the earlier name Cupressaceae s.I. by many authors, as the two families are similar in a number of morpho logical characters. Sciadopitys S. et Z., which has often been treated as a morphologically isolated member of the Taxodiaceae, has recently been considered as a monotypic family, Sciadopityaceae. The Cupressaceae s.s. may be reorganized into two subfamilies. The Cu pressoideae is composed of genera with the uppermost cone-scales infertile and can be divided into four tribes: Cnpresseae, including Cupressus, X Cupressocyparis, Charnaecyparis and Fokeinia;Thujopsideae, including Thuja, Thujopsis and Platycladusl Junipereae, including Juniperus and Microbiota; and Tetraclineae, including Calocedrus and Tetraclinis. The Callitroideae is composed of genera with the uppermost cone-scales fertile and can be divided into three tribes: Actinostrobeae, including Actinostrobus, Callitris, Fitzroya and Neocallitropsis; Widdringtoneae, including Pilgerodendron, Diselma and Widdringtonia ; Libocedreae, including Libocedrus, Papuacedrus and Austrocedrus. Five geographical distribution patterns are recognized in the 21 genera of Cupressaceae. (a) One genus, X Cupressocyparis, is a natural hybrid derived from selections in England; (b) Two genera, Cupressus and Juniperus, are distributed in Africa, Europe, Asia and North America; (c) Three genera, Thuja, Chamaecyparis, and Calocedrus, are disjnnctly distributed in Eastem Asia and North America; (d) Five genera, Actinostrobus, Callitris, Libocedrus, Papuacedrus and Widdringtonia, have limited distribution; and (e) The other 10 genera, which are monotypic, are restricted to narrow areas except Plotycladus. Three centers of genera diversity are identified in the Cupressaceae, i. e Eastern Asia with nine genera, southwestern North America with five genera, and Australia and its adjacent islands in the east　with six genera, including New Zealand,. Tasmania, New Caledonia, and New Guinea. Other important areas are western Mediterranean with three genera and Chile and Argentinawith three genera.