《中国植物志》|| 报春花科 PRIMULACEAE ||珍珠菜族 Trib. Lysimachieae Benth. et Hook. f. ||珍珠菜属 Lysimachia L.

1. 珍珠菜属——Lysimachia L.
L. Sp. Pl. 146. 1753; Klatt in Abh. Natures. Ver. Hamb. 4, 4: 1-45. 1866; Bench et Hook. f. Gen. Pl. 2: 635. 1873; Pax in Engl. et Prantl, Nat. Pflanzenfam. 4 (1): 112. 1889; Bennell et C. M. Hu in Not. Roy. Bot. Gard. Edinb. 40: 425. 1983.
直立或匍匐草本，极少亚灌木，无毛或被多细胞毛，通常有腺点。叶互生、对生或轮生，全缘。花单出腋生或排成顶生或腋生的总状花序或伞形花序；总状花序常缩短成近头状或有时复出而成圆锥花序；花萼5深裂，极少6-9裂，宿存；花冠白色或黄色，稀为淡红色或淡紫红色，辐状或钟状，5深裂，稀6-9裂，裂片在花蕾中旋转状排列；雄蕊与花冠裂片同数而对生，花丝分离或基部合生成筒，多少贴生于花冠上；花药基着或中着，顶孔开裂或纵裂；花粉粒具3孔沟，圆球形至长球形，表面近于平滑或具网状纹饰；子房球形，花柱丝状或棒状，柱头钝。蒴果卵圆形或球形，通常5瓣开裂；种子具棱角或有翅。
本属的模式种：毛黄连花 Lysimachia vulgaris L.
约180余种，主要分布于北半球温带和亚热带地区；少数种类产于非洲、拉丁美洲和大洋洲。我国有132种，1亚种和17变种，部分为民间常用草药和香料。
珍珠菜属分类系统总览
1. 香草亚属 Subgen. Idiophyton Hand. -Mazz. 花5基数，黄色，极少白色，单出腋生或排成总状花序；花丝短，长不超过花药的一半，下部合生成环，环极短，几全部与花冠基部合生；花药基着，直立，顶孔开裂，稀纵裂。植物体通常无有色腺体。
组1. 三叶组 Sect. Idiophyton 茎直立，通常仅有3枚叶聚生茎端，下部无叶。总状花序多数，在叶轮下沿茎着生；花黄色或白色。蒴果干时膜质，不开裂。
组2. 短丝组 Sect. Apodanthera Hand. -Mazz. 茎直立或匍匐。叶互生，稀2-3枚聚 生茎端。花黄色，单出腋生或排成总状花序。蒴果脆壳质，瓣裂，稀不开裂。
系1. 狭药系 Ser. Evalves Hand. -Mazz. 叶互生或聚生茎端。花药自心形基部向上长渐狭，长为基部宽的4-6倍，顶孔开裂。茎通常坚硬，多少木质化，圆柱形或具棱，很少草质而具狭翅。
系2. 宽药系 Ser. Valvatae Hand. -Mazz. 叶互生。花药多少呈钝圆锥形，长为基部宽的2-3.5倍，顶孔开裂，稀纵裂。茎草质，通常具狭翅。
组3. 对生叶组 Sect. Oppositifoliae (Hand. -Mazz.) Chen et C. M. Hu 茎草质，有狭翅，拱曲而后匍匐或鞭状伸长。叶对生。花黄色，单出或双出腋生；花药以顶端开张的 侧缝开裂。蒴果脆壳质，瓣裂。
2. 黄连花亚属 Subgen. Lysimachia 花5基数，黄色，极少白色，单出腋生或排成顶生总状花序；总状花序常缩短成头状花序状，花丝通常比花药长，下部合生成宽环或窄筒；环或筒仅基部贴生于花冠筒底部，上部与花冠分离；花药中着，稀近基着，纵裂。植物通常具有色的腺体，稀为透明腺体。
组1. 黄连花组 Sect. Lysimachia 茎粗壮，直立。叶对生或3枚轮生。总状花序复出而成圆锥花序；花黄色，萼片沿边缘有一圈黑色腺条；花丝下半部合生成筒；花药中着，纵裂。
组2. 过路黄组 Sect. Nummularia (Gilib.) Klatt 茎直立或匍匐。叶对生，稀互生或轮生。花黄色，极少（仅1种）白色，单出腋生或排总状花序或伞形花序；总状花序常缩短成近头状；花丝下部合生成宽环或窄筒；花药中着，纵裂。
系1. 浅环系 Ser. Hypericoideae (R. Knuth) Chen et C. M. Hu 花单生叶腋；花冠辐状，开张；花丝基部合生成浅环或短筒。茎直立或近直立。叶对生，除花白色者外，无柄，有透明腺点，稀具有色腺点。
系2. 侧枝系 Ser. Paracladae Chen et C. M. Hu 花在无叶的枝端或茎端排成总状花序；苞片小，与茎上部叶有明显区别；花丝下部合生成筒。茎直立或近直立。叶对生，稀轮生和互生，无柄或具有耳的柄，具有色腺点或腺条，稀无腺点。
系3. 耳柄系 Ser. Elatae Hand. -Mazz. 花单生于茎端和枝端的上部叶腋，有时稍密聚成总状花序状，稀为伞形花序；花丝下部合生成筒。茎直立。叶对生，稀轮生，披针形至狭披针形；叶柄基部通常明显扩大成耳状。植物体无腺点或具透明腺点，极少花冠具褐色腺条。
系4. 疏节系 Ser. Deltoideae Hand. -Mazz. 花单出腋生或在茎端排成疏松的总状花序；苞片叶状；花丝下部合生成筒或浅环。茎直立或上升。叶对生；叶柄具草质边缘，基部不呈耳状。植物体具透明腺点，稀具有色腺点。
系5. 叶头系 Ser. Phyllocephalae (Hand. -Mazz.) Chen et C. M. Hu 花集生于茎端或枝端成头状花序状；花丝下部合生成筒。茎直立或上升。叶对生，茎端的2-3对多少密聚，常呈轮生状。植物体不具有色腺点。
系6. 伞叶系 Ser. Paridiformes Chen et C. M. Hu 花在茎端组成伞形花序；花丝下部合生成筒。茎直立或上升。叶4至多枚轮生茎端，下部叶退化呈鳞片状。植物体有极稀疏的黑色腺条。
系7. 厚叶系 Ser. Fordianae Chen et C. M. Hu 花集生茎端成近头状或伞形花序；花丝下部合生成筒。茎直立或上升，肥厚多汁。叶对生，稍肥厚，茎端2对有时密聚呈轮生状。植物体密布黑色或褐色小腺点。
系8. 聚花系 Ser. Rubiginosae Hand. -Mazz. 花集生茎端或枝端，呈头状花序状；花丝下部合生成筒。茎匍匐或上升。叶对生。植物体具有色腺点或腺条。
系9. 过路黄系 Ser. Drymarifoliae Hand. -Mazz. 花单出腋生，极少数朵聚生茎端成近头状的总状花序；花丝下部合生成筒。茎匍匐或披散，常长蔓延。叶对生。植物体具有色腺点或腺条，稀仅具透明腺体。
系10. 金爪儿系 Ser. Grammicae Hand. -Mazz. 花单出腋生；花丝下部合生成宽而短的环。茎近直立。叶互生。植物体密生黑色腺条。
组3. 短茎组 Sect. Rosulatae R. Knuth 茎缩短，不显著或近于无茎，常自基部发出匍匐枝。叶螺旋状排列，密聚成莲座状，匙形或倒披针形。花黄色，单出腋生，具长梗；花丝比花药长，下半部合生成筒。植物体不具有色腺点。
组4. 假过路黄组 Sect. Alternifoliae R. Knuth 茎直立或匍匐上升。叶互生或有时对生。花黄色，单出腋生；花丝近分离，与花药近等长或稍短；花药近基着，直立。植物体无腺体或具有色腺点和腺条。
3. 珍珠菜亚属 Subgen. Palladia (Moench) Hand. -Mazz. 花5基数，白色、粉红色或波紫红色，排成顶生长总状花序，很少单生于茎上部叶腋或聚生于茎端；花丝分离，贴生，于花冠筒中上部或冠裂片基部；花药纵裂。
组1. 滨海组 Sect. Lubinia (Comm.) Klatt 总状花序圆锥形；苞片叶状；萼片长圆形， 边缘和顶端均膜质；花冠白色；雄蕊内藏；花药长圆形，近基着，直立；花柱钻形，长略超过花冠中部；子房圆锥形，先端渐尖。叶互生。
组2. 穗花组 Sect. Spicatae (R. Knuth) Hand. -Mazz. 总状花序多花，呈长穗状；苞片钻形；萼片椭圆形，边缘和圆钝的顶端均膜质，具腺状缘毛；花冠白色；雄蕊远短于花冠；花药卵圆形，中着；花柱棒状，长约达花冠中部。叶互生。
组3. 合萼组 Sect. Apochoris (Duby) Hand. -Mazz. 总状花序顶生；苞片钻形；花萼下部合生达全长的1/3或近1/2；花冠白色，分裂近达基部；花丝贴生于花冠裂片的近中部； 花柱长约达花冠中部。叶互生。
组4. 短柱组 Sect. Chenopodiopsis Hand. -Mazz. 总状花序疏花或花单生于茎上部叶腋而成总状花序状；苞片钻形，位于花序下部的常呈叶状；萼片披针形，边缘膜质；花冠白色，稀粉红色，通常有褐色或黑色腺条；雄蕊内藏；花药小，卵圆形，中着；花柱极短，稀长达花冠中部。叶互生。
组5. 腺药组 Sect. Miltandrae Hand. -Mazz. 花序长总状；苞片钻形，极少呈叶状；萼片披针形或钻形；花冠白色；雄蕊短于或近等长于花冠；花药卵形或线形，近基着或中着，药隔顶端具红色粗腺体或胼胝状小尖头；花柱内藏或与花冠近等长。叶对生，基部常耳状抱茎。
组6. 长蕊组 Sect. Palladia 花序长总状；苞片钻形；花冠阔钟形，极少狭钟形，白色或粉红色；雄蕊比花冠长或与之近等长；花药卵圆形或近圆形，极少卵状长圆形，中着；花柱纤细，与雄蕊等长。叶互生，很少对生或轮生。
组7. 湿生组 Sect. Candidae Hand. -Mazz. 花序总状；苞片钻形；花冠白色，狭钟形，筒部长超过全长的1/3或达1/2；雄蕊稍短于花冠；花药狭长圆形，中着；花柱自花蕾中伸出。叶互生，稀近对生。
组8. 低矮组 Sect. Pumilae (Hand. -Mazz.) Chen et C. M. Hu 花少数，集生于茎 端或单生于茎端叶腋；萼片披针形；花冠淡红色；雄蕊长于或等长于花冠；花药椭圆形，中 着；花柱约与雄蕊等长。植物体矮小，匍匐或近直立。叶互生或对生。
4. 异花珍珠菜亚属 Subgen. Heterostylandra (Hand. -Mazz.) Chen et C. M. Hu 茎花葶状，不分枝；基生叶大，莲座状丛生；茎叶少数，向上渐次缩小成苞片状；花冠淡紫色，筒状，合生部分达全长的1/2-3/4；雄蕊和花柱有长短两型；花丝下部合生成浅环或短筒。植物体不具有色的腺体。
5. 球尾花亚属 Sugen. Naumburgia (Moench) Klatt 花通常6-7基数，酪黄色；总状花序短，腋生，具短梗，因花密集而成球状或短穗状；雄蕊和花柱伸出花冠外，花丝分离。

《云南植物志》|| 报春花科 Primulaceae ||报春花科 Primulaceae ||珍珠菜属 Lysimachia Linn.

1.珍珠菜属 Lysimachia Linn.
多年生草本，少有一、二年生，极少茎下部木质化，植株呈亚灌木状。茎通常直立，有些种倾卧匍地，无毛或被多细胞柔毛，通常有细小的暗色腺点。叶互生、对生，稀轮生，全缘，两面通常有暗色腺条或腺点。花排成花序，总状、伞形或近头状，顶生或腋生，或有时单花生叶腋，稀成圆锥花序；花5数，稀6—7数；花萼深裂，宿存，常具暗色腺体；花冠黄色或白色，有时淡红或淡紫，辐状或钟状，深裂至中部或通常裂至中部以下，裂片顶端全缘；雄蕊与花冠裂片同数对生，花丝短于或显著长于花药，分离，下部贴生花冠筒部或下部合生成环或筒并贴生花冠基部，上部分离，花药基着或中着，顶孔开裂或纵裂；花粉粒具3孔沟，圆球形或椭圆体形，表面近光滑或有网状纹饰；子房球形，花柱丝状或棒状，内藏或伸出，柱头钝。蒴果球形或卵圆形，通常5瓣裂；种子具棱或有翅。
约180余种（据《中国植物志》，又据 D. J. Mabberly 1996 记载有150种），产北半球温带和亚热带，部分种产东南亚，少数种产南美、非洲、大洋洲。我国现知有136种，南、北各地均产，常见中、低海拔地区。云南有65种6变种，大部分种类见于东南部和西北部。
《云南种子植物名录》记载云南还有假露珠菜 L. circaeoides Hemsl.，异花珍珠菜 L. crispidens (Hance) Hemsl.，折瓣珍珠菜 L. reflexiloba Hand.-Mazz.，经《中国植物志》第59卷第1分册作者研究，以上3种不产云南。名录上引证的标本目前所能见到的罗开钧613499号经鉴定是阔瓣珍珠菜L. platypetala Franch.。
本属的一些种在各地民间作药用，少数种含挥发油，干植株的芳香气经久犹存。

《Flora of China》|| Primulaceae || ||珍珠菜属 Lysimachia

1. Lysimachia Linnaeus, Sp. Pl. 1: 146. 1753.

珍珠菜属 zhen zhu cai shu

Herbs erect or procumbent, rarely suffruticose, glabrous or pubescent, often with internal glands appearing as pustulate dots or stripes. Leaves alternate, opposite, or whorled, usually entire. Flowers solitary in axils of upper leaves or in terminal and axillary panicles or racemes, often shortened into capitate clusters, with bracts. Calyx usually 5(or rarely 6--9)-parted. Corolla white or yellow, rarely pink, homomorphic, rarely heteromorphic, subrotate or campanulate, deeply 5(or rarely 6--9)-parted; lobes contorted in bud. Filaments free or connate into a ring or tube at base and ± adnate to corolla tube; anthers basifixed, dorsifixed or versatile opening by apical pores or by lateral slits. Capsule subglobose, usually dehiscing by valves, rarely indehiscent.

Approximately 180 species in temperate and subtropical parts of the northern hemisphere, but with a few species in Africa, Australia, and South America; 138 species occur in China.

《Flora of North America》|| Myrsinaceae || || Lysimachia

3. Lysimachia Linnaeus, Sp. Pl. 1: 146. 1753; Gen. Pl. ed. 5, 72. 1754.

Loosestrife [Greek lysis, dissolve, and mache, strife, alluding to soothing properties]

Anita F. Cholewa

Glaux Linnaeus; Naumburgia Moench; Steironema Rafinesque

Herbs [shrubs], perennial, usually not succulent, glabrous or stipitate-glandular; resin canals sometimes obvious. Rhizomes usually present; roots fibrous. Stems erect to ascending, prostrate, decumbent, or trailing, simple or branched. Leaves cauline, opposite, whorled, or alternate, monomorphic (L. lanceolata dimorphic); petiole present or absent; blade linear to elliptic, orbiculate, ovate, lanceolate, oblanceolate, or deltate, base truncate to obtuse, rounded, or cuneate, frequently decurrent on petiole, sometimes also on stem, margins usually entire, rarely sinuate or serrulate, usually plane to revolute, glabrous, irregularly pubescent, ciliolate, or stipitate-glandular, apex obtuse to rounded, acute, or acuminate, surfaces usually glabrous or pubescent, or stipitate-glandular. Inflorescences terminal and/or axillary, racemes, panicles, [umbels], verticils, or solitary flowers; bracts present, sometimes becoming indistinguishable from leaves. Pedicels present (absent in L. maritima). Flowers: sepals 5(-9), green or whitish to pinkish, calyx deeply lobed, lobes usually lanceolate, ovate, or deltate, longer than tube; petals absent or 5(-9), corolla yellow (sometimes with reddish base) or white and sometimes darkly streaked (by resin canals) variously black, brown, maroon, red, or brownish violet, usually rotate (to funnelform or campanulate), deeply lobed, lobes longer than tube, apex variously obtuse to rounded or acute to acuminate, mucronate, or apiculate; stamens 5; filaments connate proximally or distinct; staminodes present or absent. Fruits capsular, globose, 1.3-7 mm, dehiscence valvate. Seeds 1-20+ (unknown in L. nummularia), shiny black, brown, or reddish brown, ovoid to obconic or trigonous or ± hemispheric, sometimes sharply margined or minutely winged, smooth to roughened, sometimes with deciduous, netlike, grayish covering. x = 8-15.

Species ca. 160 (20 in the flora): nearly worldwide; chiefly north temperate, especially Asia.

H. Handel-Mazzetti (1928), and later J. D. Ray (1956), divided Lysimachia into five subgenera, primarily distinguished on such characters as flower color, number of floral parts, presence/absence of staminodes, inflorescence type, etc. They also indicated that vegetative characters exhibit much variability and transitional traits (particularly in subg. Seleucia: L. ciliata, L. hybrida, L. lanceolata, L. quadriflora, L. radicans, L. tonsa). Ray recognized L. hybrida to be a subspecies of L. lanceolata and treated L. graminea as a narrow-leaved form within L. lanceolata. V. J. Coffey and S. B. Jones (1980) and most recent floras resegregate those species. Additionally, hybrids are also known to occur between some of the species (especially L. quadrifolia, L. terrestris, and L. thyrsiflora), have become well established in some areas, and contribute to taxonomic confusion. As a result, regionally, species may appear quite distinct, but in the flora as a whole, there is a blurring of some species boundaries.

Basal leaves often exhibit different states than medial and distal cauline leaves (e.g., larger blades, longer petioles, etc.). Descriptions of leaves here are based on medial and distal leaves only. Some species are described as having ciliate margins on the petiole or blade base. Petiole "cilia" frequently are actually thickened flaps of tissue, sometimes with widened bases. Flowers are pedicellate, and subtended by bracts or leaves or various transitional forms, which lead to inflorescence transitions, from solitary flowers in axils of leaves to leafy "panicles" to dense racemes. Occasionally, solitary flowers axillary to whorled leaves appear clustered, and have been referred to (as here) as verticils. Most species have a relatively short or long staminal tube. Some (subg. Seleucia) have a series of staminodes alternating, and forming a single whorl, with the fertile stamens. These staminodes are rather short and more or less lanceolate; in Lysimachia thyrsiflora, small teeth are sometimes produced on the staminal tube, perhaps vestiges of staminodes. For a more comprehensive discussion of the morphology, see J. D. Ray (1956). Clearly, more work is needed to understand the species relationships, especially in light of recent research that indicates close affinities among the species of Lysimachia and Trientalis and some Anagallis (see A. A. Anderberg et al. 2007 for further discussion).

Some species are choice ornamentals (e.g., Lysimachia ciliata, L. clethroides, L. nummularia, and L. punctata); L. clethroides and L. nummularia can become overly aggressive and even invasive.

SELECTED REFERENCES Coffey, V. J. and S. B. Jones. 1980. Biosystematics of Lysimachia section Seleucia (Primulaceae). Brittonia 32: 309-322. Handel-Mazzetti, J. 1928. A revision of the Chinese species of Lysimachia, with a new system of the whole genus. Notes Roy. Bot. Gard. Edinburgh 16: 51-121. Hao, G. et al. 2004. Molecular phylogeny of Lysimachia (Myrsinaceae) based on chloroplast trnL-F and nuclear ribosomal ITS sequences. Molec. Phylogen. Evol. 31: 323-339. Ray, J. D. 1956. The genus Lysimachia in the New World. Illinois Biol. Monogr. 24.

《Flora of China》|| Primulaceae || ||珍珠菜属 Lysimachia

1. Lysimachia Linnaeus, Sp. Pl. 1: 146. 1753.

珍珠菜属 zhen zhu cai shu

Herbs erect or procumbent, rarely suffruticose, glabrous or pubescent, often with internal glands appearing as pustulate dots or stripes. Leaves alternate, opposite, or whorled, usually entire. Flowers solitary in axils of upper leaves or in terminal and axillary panicles or racemes, often shortened into capitate clusters, with bracts. Calyx usually 5(or rarely 6--9)-parted. Corolla white or yellow, rarely pink, homomorphic, rarely heteromorphic, subrotate or campanulate, deeply 5(or rarely 6--9)-parted; lobes contorted in bud. Filaments free or connate into a ring or tube at base and ± adnate to corolla tube; anthers basifixed, dorsifixed or versatile opening by apical pores or by lateral slits. Capsule subglobose, usually dehiscing by valves, rarely indehiscent.

Approximately 180 species in temperate and subtropical parts of the northern hemisphere, but with a few species in Africa, Australia, and South America; 138 species occur in China.

《Flora of China》|| Primulaceae || ||珍珠菜属 Lysimachia

1. Lysimachia Linnaeus, Sp. Pl. 1: 146. 1753.

珍珠菜属 zhen zhu cai shu

Herbs erect or procumbent, rarely suffruticose, glabrous or pubescent, often with internal glands appearing as pustulate dots or stripes. Leaves alternate, opposite, or whorled, usually entire. Flowers solitary in axils of upper leaves or in terminal and axillary panicles or racemes, often shortened into capitate clusters, with bracts. Calyx usually 5(or rarely 6--9)-parted. Corolla white or yellow, rarely pink, homomorphic, rarely heteromorphic, subrotate or campanulate, deeply 5(or rarely 6--9)-parted; lobes contorted in bud. Filaments free or connate into a ring or tube at base and ± adnate to corolla tube; anthers basifixed, dorsifixed or versatile opening by apical pores or by lateral slits. Capsule subglobose, usually dehiscing by valves, rarely indehiscent.

Approximately 180 species in temperate and subtropical parts of the northern hemisphere, but with a few species in Africa, Australia, and South America; 138 species occur in China.

《Flora of China》|| Primulaceae || ||珍珠菜属 Lysimachia

1. Lysimachia Linnaeus, Sp. Pl. 1: 146. 1753.

珍珠菜属 zhen zhu cai shu

Herbs erect or procumbent, rarely suffruticose, glabrous or pubescent, often with internal glands appearing as pustulate dots or stripes. Leaves alternate, opposite, or whorled, usually entire. Flowers solitary in axils of upper leaves or in terminal and axillary panicles or racemes, often shortened into capitate clusters, with bracts. Calyx usually 5(or rarely 6--9)-parted. Corolla white or yellow, rarely pink, homomorphic, rarely heteromorphic, subrotate or campanulate, deeply 5(or rarely 6--9)-parted; lobes contorted in bud. Filaments free or connate into a ring or tube at base and ± adnate to corolla tube; anthers basifixed, dorsifixed or versatile opening by apical pores or by lateral slits. Capsule subglobose, usually dehiscing by valves, rarely indehiscent.

Approximately 180 species in temperate and subtropical parts of the northern hemisphere, but with a few species in Africa, Australia, and South America; 138 species occur in China.