铁线莲属研究随记(Ⅲ)

(1)在铁线莲属的15个组330余种中,只有绣球藤组sect．Cheiropsis的5个种(短梗铁线莲Clematis brevipes Rehd.,美花铁线莲C．potaninii Maxim.,绣球藤C．montana Buch．-Ham．ex DC.,薄叶铁线莲C．gracilifolia Rehd．&; Wils．和金毛铁线莲C．chrysocoma Franch．)的花与比自己原始的近缘属——银莲花属Anemone的花在构造上相近似,这表现在：萼片水平开展,膜质,倒卵形,顶端圆形或圆钝,内面无毛,外面被柔毛,但边缘无狭的短绒毛带;雄蕊无毛,花丝条形(linear)或狭条形,花药长圆形或狭长圆形,药隔顶端不突出,上述均是铁线莲属花的原始特征。在此属的其他种,可以看到萼片和雄蕊的形态发生了以下诸种变化：萼片从平展变为斜上方开展或甚至直立,质地变厚,呈纸质,有时甚至呈革质,形状多少变狭长,呈倒卵状长圆形、长圆形、倒披针形、披针形、条形,甚至狭条形,顶端有时变为渐尖或渐狭,内面出现了或疏或密的柔毛,外面边缘出现了一条极狭的短绒毛带;在雄蕊方面,花丝全长由等宽变为不等宽,呈狭披针形或狭倒披针形,花药变狭长,呈条形或狭条形,药隔顶端出现了短或长的突起,花丝或花药或二者上出现了或多或少的柔毛。上述的形态变化当均属衍生特征;(2)建立了绣球藤组sect.Cheiropsis和广布于北、南美洲和大洋洲具单性花的单性铁线莲组Sect．Aspidanthera的新分类系统;(3)描述了1新亚组,13新系,5新种,4新变种;(4)做出了6新等级,2新组合。     

铁线莲属灌木铁线莲组修订

对毛茛科铁线莲属Clematis的灌木铁线莲组sect. Fruticella进行了全面修订,确定此组共含5种、2 变种和3变型;对此组的分类学简史和地理分布做了介绍;写出本组2系的形态特征和地理分布,组下分类 群检索表,以及各种植物的形态描述、地理分布、生长环境等,并附有各分类群的3幅插图。根据花的构造 认为在铁线莲属中,灌木铁线莲组与黄花铁线莲组sect. Meclatis在亲缘关系上最为接近。二组拥有的共 同特征为: 花的4枚萼片呈黄色,斜上方开展;雄蕊花丝下部变宽,呈披针状条形。二组的区别为: 灌木铁 线莲组的萼片边缘在花开放后展宽成膜质狭翅,雄蕊无毛;而在黄花铁线莲组,萼片边缘不展宽,雄蕊花丝 被柔毛。本组5种的花构造一致;在其他营养器官形态特征方面观察到以下演化趋势: (1)叶由不分裂到羽 状全裂,由绿色到灰绿色,由狭菱状卵形到条形;(2)花序含数朵花,具花序梗和2苞片,生于当年生枝顶端, 到花数目减少到1朵,花序梗和苞片消失,花生于当年生枝的腋生短枝顶端。根据上述演化趋势认为灌木 铁线莲的模式变型C. fruticosa f. fruticosa (叶绿色,狭卵形或披针形;花序通常含数朵花,顶生于当年生枝 上)是本组的原始类型,其他种可能均源出于此类型。根据花序特征,本组被划分为2系。第1系,灌木铁线 莲系ser. Fruticosae,含4种,分布于我国黄土高原及相邻干旱地区,向北达蒙古戈壁荒漠,向南在甘肃越过 秦岭到达甘肃南部。在黄土高原北部及相邻的内蒙古西南部集中分布本系全部4种,这里是灌木铁线莲 组的分布中心。第2系,绿叶铁线莲系ser. Virides,含绿叶铁线莲C. viridis 1种,分布于我国横断山区北部的 高山灌丛草地。     

铁线莲属黄花铁线莲组修订

对毛茛科铁线莲属Clematis的黄花铁线莲组sect. Meclatis进行了全面修订, 确定此属含13种和13变种(包括1新变种和2新变种等级); 写出了此组的分类学简史和地理分布, 并对其在铁线莲属中的系统位置和组内诸种的亲缘关系进行了讨论; 还写出了此组的分种、分变种检索表, 以及各种植物的形态描述、地理分布、生长环境等, 并附有各种的插图。此组的花构造与对枝铁线莲组sect. Brachiatae的近似, 与后者在亲缘关系上相近, 区别在于此组的萼片通常斜上方开展, 呈黄色, 被毛的花丝下部变宽, 呈狭披针形, 而在对枝铁线莲组, 萼片水平开展, 呈白色, 被毛的雄蕊花丝呈狭条形, 下部不变宽; 二组可能均起源于欧洲铁线莲组的sect. Clematis subsect. Clematis, 因此, 均应是隶属欧洲铁线莲亚属subgen. Clematis的成员。根据对此组植物形态特征的分析, 观察到以下演化趋势: (1)叶的颜色由于适应干旱气候, 由绿色变为灰绿色; (2)卵形或宽卵形、掌状分裂、边缘具齿的小叶可能是原始的特征, 而披针形或条形、不分裂、全缘的小叶是衍生的特征; (3)单独、顶生、只是花梗的花是由具花序梗和二苞片的聚伞花序发生减化(reduction)而衍生的; (4)萼片形状的演化趋势与小叶形状的演化趋势近似, 也由卵形演变到披针形或条形; (5)萼片内面无毛是原始现象, 而被毛则是衍生现象; (6)萼片顶端无突起是原始现象, 出现突起则为衍生现象; (7)花药形状由长圆形演变到狭长圆形和条形。根据上述演化趋势, 推测具较多原始特征的甘川铁线莲C. akebioides和甘青铁线莲C. tangutica为此组的原始种, 而具较多衍生特征的尾尖铁线莲C. caudigera和角萼铁线莲C. corniculata为此组的进化种。组成世界屋脊的青藏高原西缘、帕米尔高原和邻近山地集中分布有此组10种(包括7特有种), 当是此组的分布中心; 而甘川铁线莲和甘青铁线莲二种分布区的主要重叠部分所在的青藏高原东缘则可能是此组的起源中心。过去,一些铁线莲属专家将属于欧洲铁线莲组的C. ispahanica Boiss.和属于对枝铁线莲组的C. graveolens Lindl.误置于黄花铁线莲组中, 对此, 本文予以纠正。     

铁线莲属一新分类系统

提出毛茛科Ranunculaceae铁线莲属Clematis一新分类系统。首先,简要回顾了此属的分类学研究历史,继对此属的营养器官和生殖器官的重要形态特征和花粉特征进行了分析,揭示出一系列演化趋势。这些趋势,尤其是萼片和雄蕊的演化趋势说明在现存的铁线莲属植物中,绣球藤Clematis montana与其少数近缘种的花构造最接近比铁线莲属原始的银莲花属Anemone,因此,包括绣球藤等植物的绣球藤组sect. Cheirpsis被认为是铁线莲属的原始群。主要根据花构造对铁线莲属现存的15组的亲缘关系进行了分析,发现它们是在4条演化干(绣球藤干、欧洲铁线莲干、尾叶铁线莲干、长瓣铁线莲干)的演化过程中先后形成的。本文将此4条演化干处理为亚属。最后做出属下各级分类群的系统排列,并给出简要的形态特征。     

中国毛茛科植物小志(廿二)

(1)揭示了铁线莲属以下演化趋势:萼片由开展到直立;雄蕊由无毛到有毛;雄蕊花丝由条形演 化到披针状条形或倒披针状条形;花药由长圆形演化到条形或狭条形;药隔不突出到在顶端突出;在雄 蕊被毛时,毛由少而短到多而长;此外花序由具花序梗和苞片到花序梗和苞片消失,以及由自当年生枝 叶腋生出转变到自老枝腋芽中生出。主要根据上述演化趋势,本文将我国铁线莲属各组及组下分类群做出新的排列。(2)描述了6新亚组,6新系,2新种,4新变种,给出了5新组合,4新等级和2新名。     

中国毛茛科植物小志(廿二)

(1)揭示了铁线莲属以下演化趋势:萼片由开展到直立;雄蕊由无毛到有毛;雄蕊花丝由条形演 化到披针状条形或倒披针状条形;花药由长圆形演化到条形或狭条形;药隔不突出到在顶端突出;在雄 蕊被毛时,毛由少而短到多而长;此外花序由具花序梗和苞片到花序梗和苞片消失,以及由自当年生枝 叶腋生出转变到自老枝腋芽中生出。主要根据上述演化趋势,本文将我国铁线莲属各组及组下分类群做出新的排列。(2)描述了6新亚组,6新系,2新种,4新变种,给出了5新组合,4新等级和2新名。     

东川铁线莲, 云南毛茛科一新种

描述了自云南东北部发现的毛茛科铁线莲属一新种,东川铁线莲。此种由于其叶为单叶,萼片直立,雄蕊的花丝和花药均被短柔毛而与曲柄铁线莲近缘,与后者的区别在于其叶呈心状卵形,叶缘具粗牙齿,花序为二歧聚伞花序,以及花药顶端钝,不具短尖头。     

铁线莲属对枝铁线莲组修订

对毛茛科铁线莲属Clematis L.的对枝铁线莲组sect. Brachiatae进行了全面修订,确定此组共含24种和4变种(包括2新种),写出了此组的分类学简史和地理分布,对其在铁线莲属中的系统位置和起源以及组下分类进行了讨论;还写出了组下分类群检索表,以及每系和每种植物的形态描述、地理分布、生长环境等,并附有多幅插图。本组各种的花构造相当一致：萼片4,镊合状排列,水平开展,白色,通常呈长圆形、卵形或披针形,只在2进化种呈宽椭圆形,外面边缘上密被短绒毛;雄蕊花丝狭条形,被柔毛,花药通常长圆形或狭长圆形,无毛,药隔顶端通常不突出;心皮密被柔毛。上述花构造与威灵仙组欧洲铁线莲亚组sect. Clematis subsect. Clematis 的花构造极为相似,与后者的区别仅在于本组被毛的雄蕊花丝。本组是在1992年由黄花铁线莲组sect. Meclatis (萼片4,通常向斜上方开展,黄色,卵形、长圆形或披针形;花丝由于下部变宽而呈狭披针形,被柔毛)中分出建立的,可能与黄花铁线莲组有亲缘关系,并可能同自威灵仙组欧洲铁线莲亚组演化而出。本组的叶变异较大,自单叶至三出复叶、1-3回羽状复叶,最后到3-4回羽状细裂,表现出一明显演化趋势。根据叶的上述特征,以及萼片形状,本组被划分为2系。第1系,对枝铁线莲系ser. Wightianae: 叶为单叶、三出复叶或1-2(-3)回羽状复叶,萼片呈长圆形至披针形,含22种,其中20种分布于科摩罗、马达加斯加、毛里求斯、非洲大陆和阿拉伯半岛西南部,另2种分别分布于印度南部和喜马拉雅山区西部。第2系,细裂铁线莲系ser. Dissectae: 叶3-4回羽状全裂,萼片呈宽椭圆形,含2种,特产马达加斯加。本组各种植物中具单叶的只有1种,即盒子草铁线莲C. actinostemmatifolia (特产科摩罗)。特产马达加斯加的伏毛铁线莲C. strigillosa和特产毛里求斯及马达加斯加的毛里求斯铁线莲C. mauritiana的叶全部为三出复叶。特产非洲大陆中部的扎伊尔铁线莲C. zaireensis的叶多数为三出复叶,有少数叶为具5枚小叶的羽状复叶;此种与产马达加斯加的伏毛铁线莲相近缘,但比后者进化,可能系由后者或后者的近缘种演化而来,并代表了非洲大陆对枝铁线莲组的原始类型,因为非洲大陆该组的其他13种均具1-2回羽状复叶。此外,特产印度南部的怀特铁线莲C. wightiana具1回羽状复叶,特产喜马拉雅山区西部的浓香铁线莲C. graveolens则具2-3回羽状复叶。根据上述,推测科摩罗、马达加斯加和毛里求斯这一群岛地区可能是对枝铁线莲组的起源中心。     

越南铁线莲属一新种

本文描述的产于越南的毛茛科Ranunculaceae铁线莲属Clematis一新种C. hagiangensis N. T. Do是欧亚大陆第一个具单性花的种, 在花构造方面与单性铁线莲组单性铁线莲亚组sect. Aspidanthera Spach subsect. Dioicae (Prantl) W. T. Wang的种类近缘, 但叶均为单叶, 萼片呈卵形或宽卵形而不同。在单性铁线莲亚组的种, 叶通常为复叶, 只在C. dimorphophylla W. T. Wang和C. variifolia W. T. Wang同时为单叶和复叶, 此外萼片呈长圆形、倒披针形或狭卵形。     

铁线莲属茴芹铁线莲组修订

对毛茛科铁线莲属Clematis L.的茴芹铁线莲组sect.Pseudanemone进行了分 类学修订,确定此组包含约16种、3亚种和2变种;写出了此组的分类学简史和地理分布,讨论 了此组在铁线莲属的系统位置;将此组划分为3系,写出了分系、分种检索表,以及各种植物 的形态描述、地理分布、生长环境等,并附有多幅插图。（2）本组植物的花具4枚平展的萼 片,雄蕊花丝条形,被柔毛。在铁线莲属中,这样的花构造与对枝铁线莲组sect.Brachiata e的花较为相似。但本组植物为直立的亚灌木或小灌木,萼片常部分镊合状排列,部分覆瓦状 排列,通常呈卵形或宽卵形,内面常密被短柔毛或短绒毛,雄蕊花药通常条形,较长,常长 达5-9.5mm。因为具有这些较为特化的特征,此组遂比对枝铁线莲组为进化（在对枝铁线莲组 ,所有的种均为木质藤本植物,萼片4枚镊合状排列,通常呈长圆形,内面常被短柔毛,但不 被短绒毛,雄蕊花药通常长圆形,较短,长1-3mm）,可能由后者演化而来,因此,茴芹铁线 莲组和对枝铁线莲组一样,也是属于铁线莲属中欧洲铁线莲演化干的一个成员。（3）如前所 述,本组的萼片常部分镊合状排列,部分覆瓦状排列。在铁线莲属中,绣球藤组sect.Cheir opsis特产日本的单型亚组subsect.Williamsianae和单性铁线莲组sect.Aspidanthera特产新 西兰的亚组subsect.Hexapetalae的萼片卷叠式也有类似情况,这说明在铁线莲属中不同演化 干上出现的部分萼片呈覆瓦状排列的情况应当是次生的,而不是原初的现象。（4）根据萼片 的毛被和花药的长度,本组被划分为3系:第1系,茴芹铁线莲系ser.Pimpinellifoliae,是本 组的原始群,含7种,特产马达加斯加中亚山区,其特征为萼片薄纸质,内面无毛或多少疏被 短柔毛,花药条形或狭长圆形,长2.2-4mm。第2系,绒毛铁线莲系ser.Villosae,含3种,特 产非洲大陆中部及南部,其特征为萼片常较厚,呈纸质或亚革质,内面密被短绒毛或密短柔 毛,花药通常条形,有时狭长圆形,长2.5-4（-5）mm。第3系,黄果铁线莲系ser.Chrysoca rpae,含6种,特产非洲大陆中部之南,其特征为萼片较厚,呈纸质或亚革质,内面通常被密 短柔毛或短绒毛,只1种无毛,花药条形,在多数种长5.8-7mm,在特产扎伊尔南部的C.kata ngensis则长8-9.5mm,是茴芹铁线莲组中最长的花药。     

A revision of Centella series Capenses (Apiaceae)

Centella capensis (L.) Domin and related species form a distinct but poorly known group within the genus. The most conspicuous diagnostic feature is the unique inflorescence structure. Each umbel comprises a central umbellule reduced to a single, sessile, functionally female flower and four lateral, functionally male, pedicellate umbellules each reduced to a single flower. The umbel is surrounded by four large, foliaceous bracts. Another unusual character is the habit. Most of the species are annuals or short-lived perennials, while all other species of the genus are perennial shrublets. A taxonomic revision of the four species and three varieties recognized, is presented.     

terminal flower: a gene affecting inflorescence development in Arabidopsis thaliana

Growth of flowering stems in wild-type Arabidopsis is indeterminate. Many flowers arise sequentially on the flanks of apical meristems in a phyllotactic spiral. We have isolated eight recessive mutants of a gene, terminal flower, in which inflorescences become determinate. Flower primordia sooner or later ‘invade’ the meristem summit leading to cessation of its further growth. Primary apical meristems usually terminate with several part-flowers which lack pedicels, and several normal pedicellate flowers may arise first. By contrast apical meristems of secondary branches usually produce only a single pedicellate flower. Plant height is also reduced and more rosette inflorescences develop. These growth patterns occur in six strong mutants raised at 25°C under continuous light. In two weak mutants termination occurs much later with many more flowers arising before eventual termination. Termination is similarly delayed in at least one of the strong mutants grown at lower temperatures. The tfl mutation does not affect the indeterminate growth of flower meristems, at least in-so-far as this occurs in agamous mutants. The tfl locus is at the top of linkage group 5, close to RFLP 447. We propose that the TFL gene product supports the activity of an inhibitor of flower primordium initiation. This inhibitor would normally prevent flowers from arising on the inflorescence apex but in tfl mutants it may readily fall below its threshold of activity. The TFL gene may be one of a class responsible for evolutionary changes between indeterminate and determinate growth.     

Honeybee (Apis mellifera ligustica) Response to Differences in Handling Time, Rewards and Flower Colours

Free flying honeybees were tested outdoors on blue–white and blue–yellow dimorphic artificial flower patches to examine the influence of reward difference, flower handling‐time difference and flower colour choice on foraging decisions. We employed different flower‐well depths to vary handling times (costs), and differences in sucrose molarity to vary reward quality. Tests were performed with 2 and 6 μl rewards to vary quantity. We show that when handling time is correlated with flower‐colour morphs on a pedicellate artificial flower patch, a honeybee's foraging behaviour is dependent on the flower colours used in the choice tests. This supports a honeybee foraging model where constraints are a significant factor in decision making. Bees visiting blue–yellow flower patches exhibited flower constancy to colour, where they restricted most visits to a single flower colour, some bees to blue and others to yellow, irrespective of handing time differences. When offered a choice of equally rewarding blue or white flowers, bees were not constrained by flower colour and chose to visit flowers with a lower handling time. When reward molarity varied with well depth between blue and white flowers, foragers chose shallow‐well flowers (short‐handling time) with a smaller net harvest rate over deep‐well flowers (long‐handling time) with a greater net harvest rate. Results using the blue–white dimorphic flower patch suggest that when foraging options simultaneously involve reward and handling‐time choices, honeybee forager behaviour is inconsistent with an absolute method of evaluating profit.     

On the morphological position of Paepalanthus subgenus Psilandra (Eriocaulaceae)

Paepalanthus subgen. Psilandra comprises two species known only from the type collections. The aim of this study is to evaluate their unique floral morphology within a given phylogenetic context and its implications for taxonomy. Flowers of both species were analyzed, described, and photographed using a digital microscope. Taxonomically relevant characters were mapped on previous molecular phylogenies. The flowers of both species fit the Paepalanthoideae pattern and are quite similar to each other. The petals of the pistillate flowers, although free, show evidence of previous fusion in the middle. The free condition is interpreted to be secondary, probably because of the handling of the specimens or to the current developmental stage. Character mapping indicates that both species share floral features with Syngonanthus but not with Paepalanthus. Therefore, based on the median fusion of the pistillate flower petals, absence of floral bracts, petals of the pistillate flower being shorter than the sepals, simple stigmas in the commissural position, the pedicellate pistillate flower, and the glabrous petals and sepals, we transfer P. subgen. Psilandra to synonymy of Syngonanthus sect. Syngonanthus and combine both species in this genus.     

An extinct genus of Salicaceae based on twigs with attached flowers, fruits, and foliage from the Eocene Green River Formation of Utah and Colorado, USA

A newly recovered twig with attached leaves and flowers from the Eocene Green River Formation of Utah provides the basis for recognizing a new, extinct genus of Salicaceae sensu lato (s.l.). Pseudosalix handleyi gen. et sp. nov. has alternate lanceolate leaves with pinnate, semicraspedodromous venation and a serrate margin with glandular teeth. The inflorescence is terminal on the twig and is unisexual, composed of flowers organized in a paniculoid cyme, with lateral paraclades of pedicellate flowers. The attached pistillate flowers have four prominent sepals that are valvate in bud, spreading but basally fused at anthesis; the single pistil of each flower is ovoid with three or four longitudinal sutures, indicating development to a capsular fruit. Three or four recurved styles radiate from the apex of the pistil, each with a distal globose stigma. The infructescence, verified by attachment to twigs with the same kind of leaves, bore capsular fruits of three and four valves. Associated but unattached, staminate flowers also have four well-developed, basally connate sepals. They are pedicellate and bear several stamens, each with a short filament and globose anther. The available morphological characters place the fossil species within the Salicaceae s.l. as an immediate sister to the clade containing Populus and Salix. Although the likely outgroup genera (including Itoa, Poliothyrsis, Carrierea, and Idesia) to tribe Saliceae all occur in Asia today and not North America, the occurrence of both Pseudosalix and Populus in the Eocene of Utah raises the possibility of a North American origin for the Saliceae.     

Primary teeth in Anura are nonpedicellate and bladed

1. Scanning electron microscop of the dentition of four anuran species (Discoglossus pictus, Bombina orientalis, Rana cyanophlyctis, Rana temporaria) revealed that at least the primary teeth, which are established during metamorphosis, are more or less bicuspid, bladed and nonpedicel-late. Juveniles and adults, however, have pedicellate teeth. 2. Thus, regardin the pedicellate condition, the sequence of stages in tooth development of Anura und Urodela show some similarities, although the primary teeth of urodeles develop already before metamorphosis.     

Astragalus sect. Astragalus (Fabaceae) in Iran, complementary notes with a key to the species

Astragalus zarreianus. is described and illustrated from Iran. It belongs to section Astragalus. This new species confined to western part of Iran (Prov. Illam) from only one location. A key to the species of the section known from Iran is prepared. Differences between the section and the closely related sections are discussed. On the basis of some important trends, an informal grouping is undertaken for yellow flower Astragali. Moreover, A. pseudo-orthocarpus is considered as a valid species in this treatment and differences with its closest relatives are described.     

Aloe nugalensis sp. nov. (Asphodelaceae), a new gypsum endemic from northeastern Somalia

The new species Aloe nugalensis Thulin, a shrubby plant with long, drooping leaves and a drooping, branched inflorescence with long‐pedicellate orange red flowers, is described from a gypsum hill in the Nugaal valley of northeastern Somalia. The only known material is an individual grown in the Botanical Garden of Uppsala University from seeds collected in Nov 1985 that first flowered in Feb 2011.     

The altitudinal and geographical distributions of flower types in Rhododendron section Vireya, especially in the Papuasian species, and the& significance

Most species of Rhododendron section Vireya from Papuasia can be assigned to well-marked flower types, 127 species being placed in five of these types. It is suggested that two of the main types may be pollinated mainly by birds, one by sphingid moths and one by butterflies. Above 3000 m red-flowered species greatly predominate over those with other colours in Papuasia, and also through much of Malesia; the relationship between ornithophily and this correlation is discussed. The distribution of flower types within supraspecific taxa suggests that some rearrangement of the latter may be necessary; some flower types are demonstrably polyphyletic. It is shown that in west Malesia species with flower colour other than red, not or slightly zygomorphic flowers and relatively longer corolla lobes are commoner than within Papuasia; it appears that section Viveya is, florally at least, most specialized in this latter area. Flower type and pollinators in section Vireya are related to those in the rest of the genus, and it is suggested that the adoption of ornithophily has been an important factor in the success of the Ericaceae as a whole in tropical montane areas.