作者：黄尔峰、王晖、贾宏炎
出版社：辽宁科技出版社
格式: AZW3, DOCX, EPUB, MOBI, PDF, TXT
球兰：资源与栽培试读：
前言
Foreword在我国，以兰（ Lan）为名的植物很多，它们中的大部分属于兰科Orchidaceae，但也有一些例外，如木兰 Magnolia 、马兰 Aster indicus 、金粟兰 Chloranthus spicatus 等，球兰 Hoya carnosa也是其中的一种。东汉的《说文解字》对“兰”的解释是具有香气的草本植物，球兰的花序形似绣球，花芳香，所以这一名称对它的形态和气味作了很好的概括。在 1959 年出版的《广州植物志》中记录了球兰和凹叶球兰 Hoya kerrii 2 个种，并提及它们在广州均是常见的观赏植物。该书的主编是著名植物学家，中国科学院院士陈焕镛先生，1984 年李秉滔教授发表的椰香球兰 Hoya chunii 即以他的姓氏命名。现在球兰一名不仅指 Hoya carnosa 这个种，更多的代表了球兰属Hoya 这个类群的植物。球兰属是夹竹桃科 Apocynaceae 中较大的一个属，有 300 多种。它们的叶片、花序和花的形态变化很大，观赏价值高，是优良的园艺植物。 1810 年植物学家罗伯特 . 布朗（Robert Brown）建立球兰属时，便以园艺家托马斯 . 霍伊（ThomasHoy）的姓氏命名，似乎预见了球兰属植物将会成为园艺界的宠儿。
In China, many of the Chinese plant’s name possess ‘Lan’（兰）, in which, most of them are the member of orchid family (Orchidaceae) and few of them are not having, such as ‘Mu Lan’ (Magnolia ), ‘Ma Lan’ (Aster indicus ), ‘Jin Su Lan’ (Chloranthus spicatus ) and so on. ‘Qiu Lan’ (Hoya carnosa ) is also a plant name along with ‘Lan’ in its Chinese name, which belongs to the dogbane family (Apocynaceae).The explanation of Chinese character “Lan” （ 兰）is present in the book ‘Origin of Chinese Characters ’ which was written in the Eastern Han Dynasty (121 AD). In this book, Chinese’s character was explained this word ( 兰) as a kind of herbaceous plant withfragrance. In this case, the name ‘Qiu Lan’ is also summarized well about morphology and smell: ball-shaped (in Chinese ‘Qiu’ means ball or ball shaped) inflorescence and fragrant flowers. There were two Hoya species recorded in ‘Flora of Guangzhou ’ (published in 1959): Hoya carnosa and Hoya kerrii , both of them were common ornamental plants in Guangzhou City. The chief editor of this book was the famous botanist Chun Woon-Young, an academician of Chinese Academy of Sciences. Later, Hoya chunii was described by Professor Li Ping-Tao in 1984. Now, the Chinese name ‘Qiu Lan’ refers not only to Hoya carnosa , but also to the genus of Hoya . Hoya is one of the prevalent genera of dogbane family (Apocynaceae), which contains more than 300 species. Leaves, inflorescence and flowers are various in shapes and beautiful. In very particular, these characters make them as an excellent ornamental resource in the horticultural world. In 1810, Robert Brown established the Hoya genus and named them after his friend, Thomas Hoy, a horticulturist. His contribution might be the botanist’s foresight that Hoya plants would become the favorite of horticultural world.
本书的第一作者自 1995 年开始从事球兰属植物的引种与栽培工作。初期由于缺乏参考资料，在种类鉴定和栽培养护上都经过了一番摸索。后来随着经验的积累和设施的改善，收集的种类逐步增加。同时，在国内可供栽培的球兰种类也越来越丰富，估算约有 300 种及栽培品种的球兰属植物栽培在各地的植物园、苗圃和爱好者家中。专门介绍球兰属植物引种和栽培的网站—“中国球兰网 ”（ www.hoyabbs.com）也为球兰爱好者提供了一个交流的平台。但由于这些球兰属植物大多来自国外，在国内依然缺少系统的介绍，这对它们的栽培、养护、推广和应用，以及栽培者的经验交流和分享均有诸多不便。为此，我们根据积累的资料，选出国内常见栽培的球兰属植物120 种、 1 变形和 11 个栽培品种编撰成此书，并挑选了 1200 余张彩色照片展示这些球兰的宏观形态特征，读者可以通过这些了解球兰属植物的多样性。同时，本书对这些球兰的栽培习性作了介绍，以供栽培者参考。
The author of this book began to introduce and growing Hoya plantssince 1995. Due to lack of references and experience at early stage, the cultivating work has been carried out very slowly. With an accumulation of experience and adequate facility, the collection was gradually increased. At present, an enormous Hoya species is now reproduced and available for the plant growers in China.Horticulturists are cultivating nearly 300 species of Hoya species in botanical gardens, plant nurseries as well the amateurs’ home.In the meantime, the website (www.hoyabbs.com) also provides a place for the communication of Hoya amateurs. However, the information of Hoya plant’s resource and cultivation are still very limited in China. Consequently, with accumulated information besides photos, we described 120 Hoya species, and most of them are widely-cultivated in China. Our present book also includes morethan 1200 photos in order to provide each plant’s morphological features. We hope that the readers can have a better understanding of Hoya species diversity from this book; in addition, it provides the cultivation information of each plant.
华南农业大学的李秉滔教授在此书的编写过程中给予了全面的指导，李教授是夹竹桃科的分类专家，对球兰属植物的形态、分布和分类历史非常熟悉。他不仅为本书作序，还审阅了书中所有的文字内容，在此向李教授表示衷心的感谢。本书的编研得到了深圳市城市管理局科研项目（项目编号： 201415）及中国林业科学研究院热带林业实验中心主持的科研项目（项目编号： 2005DIB6J144；SFA2130218）的支持。仙湖植物园的邱志敬博士、黄义钧、郎校安和梁文超在种类收集和栽培养护上给予了帮助， Baskaran Xvier Ravi 博士帮助审阅了本书的英文部分，林漫华和郑曼枞帮助整理稿件和名录；Rungson Phomchareon 先生在引种和照片收集工作上给予了很大帮助，作者在此深表谢意。
We are indebted to our honorable Professor Li Ping-Tao, who has given a comprehensive guidance for species collection, identification and reference collection. We thank the Shenzhen Urban Management Bureau (Project Number: 201415) and Experimental Center of Tropical Forestry, Chinese Academy of Forestry (Project Number: 2005DIB6J144 & SFA2130218) for financial and field work assistance. Many people collaborated and contributed with full of dedication in the preparation of this book,Dr. Qiu Zhi-Jing, Mr. Huang Yi-Jun, Mr. Lang Xiao-An and Mr. Liang Wen-Chao was helped for species collection and plant’s cultivation.Our heartfelt thanks to postdoctoral fellow, Dr. Baskaran Xavier Ravi helped to review and preparation of this book. Ms. Lin Manhua and Ms. Zheng Man-Cong were kindly helped in sorting and arrangement of the name list. Authors are cordially thanks to Mr.Rungson Phomchareon for providing gorgeous photographs.
本书仅收集了常见栽培的球兰属植物 120 种， 1 变形和 11 个栽培品种，但随着引种工作的深入和爱好者的增加，将来会有更多的球兰属植物应用在园林和园艺方面。我们也在继续补充收集资料，以期向读者展示更多的种类。由于编写仓促，书中难免有错误和不足，恳请读者批评指正。
Most species of Hoya species in this book are widely-cultivated in China. By introduction of Hoya species and their reproduction, as well amateur’s interest, the number are gradually increasing in China. We are still collecting information and photos in order to represent more Hoya species. Due to limited schedule, there might be mistakes in this book and criticism from readers is also welcome.编者2015 年 7 月于深圳The authorsJuly 2015 in Shenzhen City
绪言
Introduction球兰属（Hoya ） 隶属于夹竹桃科（Apocynaceae） 萝藦亚科（Asclepiadoideae ），有300 余种，分布于亚洲至大洋洲及太平洋岛屿的热带、亚热带地区。球兰属植物多为附生的藤本，茎柔软，缠绕或依靠不定根攀援于乔木或岩石上，稀为附生或地生的半灌木；叶通常对生，有时其中一片退化（如覆叶球兰Hoya imbricata ）；该属植物的叶片形态多样，通常革质或肉质，少数种类叶片较薄，表面光滑或粗糙，有时被毛，叶片通常为绿色，有时因生长环境使得一些种类的叶片变为红色、橙红色或紫红色，也有一些种类的叶片表面具有银色、银灰色或橙红色的斑点或斑块。球兰属植物的花序是其区别于其它萝藦亚科植物的主要特征之一，多数种类的花序梗在花或果实脱落后并不随花和果一同脱落，并在下一花期可继续开花。对这种特殊的花序目前还没有统一的定义，常被描述成伞形花序（Umbel）、假伞形花序（Pseudumbel）或伞形状的花序（Umbel-likeinflorescences），为便于理解，本书根据其外形统一使用伞形花序（Umbel）描述。与其它萝藦亚科的植物一样，球兰属植物花的构造复杂而精巧，是有花植物中花形态最复杂的类群之一。其花冠辐射对称，有不同的形态和颜色，花冠之上是五角星形的副花冠，通常大而明显，副花冠裂片间为导轨，花粉块藏于其中，而副花冠中心则是柱头，常被雄蕊的附属物覆盖。球兰属植物的雌蕊虽然具2 个子房，但其中一个通常不发育，所以果为单生的蓇葖果，即一个果梗上仅具一枚果，而不像其它一些萝藦亚科的植物在一个果梗上具一对蓇葖果（果双生）。
Hoya genus belongs within the dogbane family (Apocynaceae) and milkweed subfamily (Asclepiadoideae ), which consists of more than 300 species that native to the tropical and subtropical regions of Asia, Oceania and Pacific islands. Most of them are epiphytic herbaceous climbers and their soft stem twine around trees, rocks or climb on a substrate with adventitious roots.Only few of them are epiphytic or terrestrial subshrubs. Almost all species produce two opposite leaves at each node and sometimes with one of the two opposite leaves degenerating (eg. Hoya imbricata ). In general, leaf shape of different species is extremely variable but usually leathery or fleshy, and very few are thin. The roughness, degree of hairiness and color of leaf are also greatly varied in different species. Moreover, some species have silver, silver gray or orange spots or patches over the leaf surface. The inflorescence of Hoya is one of its main features that make it different from other Asclepiadoideae plants. Peduncle of most species does not fall off with flowers or fruits, and instead, that can continue to bloom in the next florescence. A special type of inflorescence present in Hoya genus, which is often described as umbel, pseudumbel, or umbel-like inflorescences. In this book, umbel is used to describe the inflorescences of Hoya according to its appearance. Like other Asclepiadoideae plants, the flowers of Hoya are mostly elaborate, complicated and delicate among flowers of angiosperms.In addition, their actinomorphic corolla has various shapes and colors. Staminal corona consists of five erect lobes, usually large and distinct. There are guide rails between corona lobes, in which pollinia hides. Stigma situates in the center of corona and usually covered by anther appendage. Hoya species have two ovaries, however, one of which usually grows out into the follicle. Therefore, the fruits are solitary follicle - one fruit on each pedicel, unlike some other Asclepiadoideae plants, which have a pair of follicle (binate fruits) on each pedicel.
在以往的分类系统中，球兰属为萝藦科（Asclepiadaceae）的一个属。但根据近期植物系统分类学的研究，整个萝藦科应作为夹竹桃科下的一个亚科，本书亦采用此分类观点，将球兰属置于夹竹桃科之下。
In previous classification system, Hoya genus belongs to milkweed family (Asclepiadaceae). According to recent research on plant systematic, Asclepiadaceae is recognized as a subfamily of dogbane family (Apocynaceae). In this book, we have adopted the same opinion that genus Hoya is belonged to Apocynaceae.球兰属植物的分类 Classification
球兰属Hoya R. Br. 由英国著名植物学家Robert Brown 于1810建立，其属名Hoya 是为纪念他的好友，英国园艺家Thomas Hoy（图1）。自球兰属建立至今，已陆续发表了600 余种，仅2014 年就发表了30 余个新种，这其中虽然存在了大量相同物种被重复发表的问题，但也说明了还有很多种类未被发现。1885 年，英国植物学家Joseph Dalton Hooker 在他编写的英属印度植物志（The Flora ofBritish India）中，对收录的球兰属植物进行了分类，这是球兰属植物的第一个较完整的分类系统（Hooker，1883）。此后，许多植物学家均尝试对球兰属植物进行分类修订。但直至现在，该属的分类问题仍然困扰着植物学家，加上每年不断有新种的发表，所以目前球兰属还没有一个完备的分类系统。而对于全世界有多少种球兰，也没有确切的数字，现在被普遍接受的为300 种左右。
Hoya genuswas named in 1810 by the prominent British botanist, Robert Brown to honor to his friend, the British horticulturist Thomas Hoy (Figure 1). Presently, more than 600 species described; of them, over 30 new species were recently described in 2014. Even though some of them might be repetitively described, yet there are many undiscovered species. In 1885, British botanist Joseph Dalton Hooker proposed a classification of genus Hoya on ‘The Flora of British India’ which is the first more complete classification system of the genus (Hooker, 1883). Later on, many botanists had tried for the comprehensive taxonomic revision of Hoya , but so far it hasn’t yet been attempted. Moreover, so many new species are continually described every year. Therefore, both classification systems of the genus and delimitation of species within the genus are still unsatisfactory.
图1.1810 年Robert Brown 在新荷兰植物长篇志（Prodromus Florae NovaeHollandiae）中建立了球兰属Hoya （文献图像来源于http://www.botanicus.org）
Figure 1. In 1810, Robert Brown established genus Hoya in ‘Prodromus Florae Novae Hollandiae’ . (Image was from http://www.botanicus.org)球兰属植物的分布与生态 Distribution and Ecology
球兰属植物的分布
Distribution
1810 年Robert Brown 在新荷兰植物长篇志（Prodromus FloraeNovae Hollandiae）中建立了球兰属，新荷兰是澳大利亚的旧称，为球兰属的自然分布区域。该属是典型的热带亚洲至大洋洲分布的属，分布范围为亚洲至大洋洲的热带、亚热带地区。其分布区的北缘位于我国西藏的南部及东南部，向南可到达澳大利亚的东北部，西缘位于印度南部的西高止山脉附近，向东则可到达南半球的斐济群岛（图2）。球兰属植物的分布与亚洲及大洋洲的热带雨林和热带季雨林的分布有很高的一致性，大部分的球兰属植物生长在热带雨林和热带季雨林的林缘地带，如森林中的河边和路边，甚至在海岸边，它们通常附生在乔木或灌木的茎干或枝条上，但也有少数较耐寒的种类生长在亚热带的常绿阔叶林中，如我国的贵州、云南等地的常绿阔叶林下。位于赤道附近的苏门答腊岛、婆罗洲岛、苏拉威西岛和新几内亚岛东部是球兰属植物的分布中心，这片区域中球兰的种类最丰富。在垂直分布上，球兰属植物多生长在海拔1000m 以下的区域，但也有部分的种类可以生长在海拔近2000m 的地方，如婆罗洲岛和新几内亚岛的高山上均有球兰的分布。
In 1810, Robert Brown described Hoya in ‘Prodromus Florae Novae Hollandiae’ . Novae Hollandiae is the former name of Australia, which is the natural distribution area of Hoya species. Genus Hoya is typically distributing in tropical and subtropical regions of Asia and Oceania continents. This distribution area ranges north to the west and southeast of Tibetan highlands, south to the northeast of Australia, west to the Western Ghats of southern India, and east to Fiji Islands (Figure 2) which corresponds rather accurately with the distribution of rainforests and monsoon forests in Asia and Oceania continents. Most Hoya species grow on the forest edges intropical area, such as river banks, roadsides within forest and even shores, which usually a climb on the stems or branches of trees and shrubs.A few cold-resistant species can be found in evergreen broadleaf forests in subtropical regions, such as Guizhou and Yunnan of China, etc. The core area of Hoya species distributed in Sumatra Island, Borneo Island, Sulawesi Island and eastern New Guinea Island, in addition, a variety of species can be found in these near equatorial regions. About the vertical distribution, most Hoya species grow below 1000 m, and only a small number of species can be found at a higher altitude about 2000 m, like mountains of Borneo Island or New Guinea Island.
图2. 球兰属植物的地理分布（根据Molecular Phylogenetics and Evolution39: 723, fig.1. 2006. 仿绘）
Figure 2.Geographical distribution of Hoya (redraw from Molecular Phylogenetics and Evolution, 39: 723, fig.1. 2006. )
球兰属植物的生态
Ecology
大部分球兰属植物是热带雨林中的附生植物，通常生长在雨林的林缘地带。它们利用茎和不定根攀附在乔木或灌木的树干上，也有部分种类附生在岩石上（主要为石灰岩）。无论在形态结构、生理功能还是繁殖方式上，球兰属植物都具有热带附生植物的特点，从而与栖息地的生态环境相适应。
Most Hoya species are epiphytic plants living in tropical rainforests, typically on the edges of forests. They climb on the branches of trees and shrubs with their soft stems and adventitious roots, while some species also grow on, usually, calcareous rocks. To adapt this habitat, the morphology, physiological functions and reproductive pattern of Hoya plants entirely feature tropical epiphytic plants.
球兰属植物多为攀援或悬垂藤本。除少数的地生种类外，植株一般具有发达的气生根（不定根），这些气生根除了具有吸收功能外，还有很强的附着能力。植株一般依靠这些气生根攀附在乔木的主干上。发达的气生根还为球兰属植物的无性繁殖提供了便利，当较长枝条折断后，断枝上的气生根可以维持水分供给，从而生长成独立的植株，这一特点让多数球兰属植物的扦插繁殖均有较高的成活率。但也因气生的原因，根需要很好的透气性，如栽培基质紧密或积水导致透气性下降则根部很容易腐烂。
Most Hoya plants are climber or hanging vines except some terrestrial species, and generally, they have well-developed adventitious roots. This kind of root is well in absorption and adhesion, by which the plants can also climb the trunk of trees.Developed adventitious roots also offer great convenience for Hoya plant’s asexual reproduction. When a long branch breaks off, the adventitious roots on this cutting will continuously provide water supply to grow into a new plant. The propagation of most Hoya plants has a high survival rate. However, the roots are aerial, and they need substrates of high air permeability. If the substrate is compact or water-logging, the roots may rot quickly.
球兰属植物的茎通常较柔软，木质化程度不高。根据茎的攀附能力，大部分的球兰属植物可分为攀援型和悬垂型两类。攀援型的球兰的着生位置通常较低，如乔木的茎干基部或中部，并通过茎和气生根在乔木的茎干上向上攀援。悬垂型的球兰着生的位置通常较高，如乔木的侧枝上。它们的茎通常不攀援，而是向下悬垂在空中。两种类型的球兰均充分利用了雨林中林冠层以下，灌木层以上相对空旷的空间作为栖息地。少部分球兰属植物的茎木质化程度较高，它们的茎既不攀援也不悬垂，而是直立或斜展，呈半灌木状（与茎完全木质化的灌木相区别），这些种类的球兰大部分也附生于林下的树干上或石上，仅有很少的种类为地生植物，如蜂出巢Hoya multiflora。
The stems of Hoya plants are herbaceous and soft in nature. Mostly, Hoya plants can be classified into climber or hanging vines.Climbers usually grow in lower position, such as the base or central part of tree trunks, on which the vines climb upward by stems and adventitious roots. Hanging vines grows in higher position, such as branches of trees. The stems of hanging vines usually hang on the air. Both two types inhabit in the empty space between forest canopy and shrub layer. Very few Hoya species having medium woody stems, which are not climbing or hanging, but erect or spread, subshrub-like (different from shrubs with entirely lignified stems), and mostly grows on rocks or trunks. Only very limited number of Hoya species are terrestrial plants, such as Hoya multiflora.
球兰属植物的叶通常呈肉质（图3），这也是园艺学家常将其归入多肉植物的原因。这种肉质的叶片与多数多肉植物一样，可以通过景天酸代谢途径（Crassulacean acid metabolism，CAM）在夜间吸收二氧化碳，并储存在叶肉细胞中，肥厚的叶片是二氧化碳的容器。具有这种二氧化碳固定机制的植物通常生长在高温、干旱的环境下，这与球兰属植物生长的雨林环境有很大的差别。但从植株生长的小环境来看，CAM 途径对球兰属植物的生长具有一定的生态学意义。附生的球兰仅能从其附着的树干或岩石上获取水分，这类基质的保水能力差，干湿变化大，一旦干燥时间过长植株就将面临缺水的胁迫，影响正常生长。在此胁迫下，球兰属植物可以利用CAM 途径在蒸发量较小的夜间吸收二氧化碳，而在蒸发量大的日间则关闭气孔，同时利用储存的二氧化碳进行正常的光合作用，维持植株的生长。而地生的球兰，如蜂出巢Hoya multiflora 和爪哇球兰Hoya javanica ，由于水分供给相对稳定，并不具有肉质叶这一特征（图4）。
Hoya species usually have succulent leaves (Figure 3). Most of them are considered as succulent plants by horticulturalists. In general, most succulent plant’s fleshy leaf absorbs CO2 through Crassulacean acid metabolism (CAM) at night and stores them in the mesophyll cells. The plants are able to store CO2 often grow in high-temperature and droughty environment, which is totally different from rainforest environment. However, CAM pathway is necessary for Hoya plants because they can only absorb water from the trunks or rocks surface. Due to low water retention capacity, moisture of substrate changes greatly. Once the substrate stays in dry condition for a prolonged period, the growth of plants might suffer from drought. In such case, Hoya plants can absorb CO2 through CAM at night during the evaporation capacity is rather low. Meanwhile, the plants use the stored CO2 for photosynthesis in a daytime while the evaporation capacity is high. For terrestrial Hoya species, such as Hoya multiflora and Hoya Javanica , due to the relatively stable water supply, the leaves are usually thin (Figure 4).
图3. 凹叶球兰Hoya kerrii 的肉质叶片
Figure 3. Fleshy leaves of Hoya kerrii
图 4. 爪哇球兰Hoya javanica 的叶片
Figure 4. Thin leaves of Hoya javanica
所有球兰属植物均为虫媒植物（依靠昆虫传粉），并且目前已知的种类中除了南方球兰Hoya australis 由昼行性的一种蝴蝶协助传粉外，其余的种类均由夜行性昆虫协助传粉，主要是蛾类和甲虫类，可能还包含了部分竹节虫类。常与球兰属植物共生的蚂蚁可能也是其传粉者之一，但所起的作用不大。球兰属植物的花具有吸引夜行性昆虫的特点，它们的花通常在傍晚开放，多数种类的花冠为白色、粉红色、黄绿色或淡黄色，这些颜色在夜晚均较显眼（图5），并且花在夜间产生蜜露和香气。根据观测（Altenburger&Matile，1988），球兰Hoya carnosa 的花释放香气具有很强的节律性，夜间香气浓而白天较淡，中间间隔为12 小时。球兰属植物花的结构精巧，具有高效的虫媒传粉机制。它们的花粉并不像大多数种子植物的花粉呈分散状，而是聚集成块，称为花粉块（Pollinium），2 枚花粉块与其下的花粉块柄（Caudicle）及着粉腺（Retinaculum）共同组成联合体，称为花粉器（Pollinarium）。每枚雄蕊产生一枚花粉器，并藏于副花冠间的导轨（Guide rail）中。当传粉者停留在花上时，足部容易踩入导轨并带出具粘性的花粉器，从而帮助传粉，这一传粉机制与兰科植物的非常相似。
All Hoya species are an entomophilous plant (pollinated by insects). Hoya australis , which is diurnally pollinated by a day flying butterfly, while the pollinator of all Hoya species could be nocturnal insects, moths, beetles, mainly and probably walkingsticks included too. Ants are live in associate with Hoya species that might be a poor pollinator. The flowers of most Hoya species open at dusk which combined various flower colors such as white, pink, yellowish green and light yellow (Figure 5). These conspicuous flowers usually produce with a high amount of nectar and strong fragrance, which main play as attractants for the nocturnal insects. By experimental(A ltenburger&Matile, 1988), flowers of Hoya carnosa emitted fragrance in a circadian rhythm which is strong in dark period and light in daytime, in other words, flowers timed their emission of fragrance to occur 12 hours after a last emission period. Flowers of Hoya species are delicate with efficient pollination mechanism. Unlike most seed plant’s disparate pollen, Hoya species have cloddy pollen, which also known as pollinium. Each anther produces two pollinia that attached through a caudicle to the retinaculum to form a pollinarium. Pollinatorsbring out the sticky pollinarium while visiting the flower in order to proceed pollination. Moreover, the mechanisms of such a case are similar to many Orchidaceae species.
图5. 球兰Hoya carnosa 白色的花冠在夜晚很显眼
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