摘 要:本文报道我国北部药用甘草的生态学特性和群落分类。目的是为人工驯化栽培提供生态学依据,并为甘草野生资源的利用和保护提供资料。
关键词:药用甘草;生态学特性;群落分类
药用甘草主要指《中华人民共和国药典》所规定的3种:甘草(Glycyrrhiza uralensis Fisch.ex DC.)、光果甘草(G.glabra L.)和胀果甘草(G.inflata Bat.)。我国药用甘草主要分布于北纬37°~48°,东经75°~132°,由西向东即由新疆的喀什至黑龙江的佳木斯,由北向南即由新疆的阿勒泰和黑龙江的龙江至新疆的和田和陕西的文县。
依生态因子分析,药用甘草均产于干旱、半干旱地区,但甘草并非旱生植物,其生境气候湿润度在1.0~0.12之间,即湿润和半湿润、半干燥地区,如果分布于湿润度<0.12的干燥环境条件者,其小生境是干燥地区的河流沿岸或地下水位较高。甘草产地年均降雨量一般在100~389mm,有些产地年均降雨量<100mm,但多为河滩、河沿岸或地下水位较高的环境,例如:内蒙古的额济纳旗,甘肃的敦煌、酒泉、张掖,青海的柴达木盆地的德令哈、都兰垦区,新疆塔里木盆地周围的于田、若羌、且未、焉耆、库车、吐鲁番等地。产地大气年均相对湿度在37%~73%之间。药用甘草主要分布于寒温带、中温带和暖温带3个温度带。年平均温度在0.4~14℃之间,最低限为阿勒泰,最高限是吐鲁番,其他产地年均温度一般为3~12℃之间。产地年积温(≥土0℃)在2 000~5 500℃的幅度,最低不低于1 900℃,如阿勒泰的北屯垦区。药用甘草产地群落分类采用三级分类单位,即高级单位植被型(Vegetation type),中级单位群系组(Formation-group)及群系(Formation)以及基本单位群丛(Association)。植被型在外貌、层片结构、生存外界条件的适应方式一致,它由若干群系组组成;群系组具有相同的生活型及生存于比较一致的环境条件,层片结构与科属组成均相似;群系的优势层建群种完全一致或具有共同建群种的所有群丛,它的生物生产力及动态趋势基本相似;群丛相当于分类学中的种(species),它在优势种和生态指示种的成分和数量关系相似,层片结构和季节变化幅度一致,对环境条件的效应及演替阶段相同,它是由若干能流和物流转化相同的群落组成。
根据以上原则,我们把甘草属植物的群落归纳为4个植被型,13个群系组,31个群系,81个群丛(不包括农作物群系组的群丛)。
1 森林植被型(Drymion)
1.1 荒漠河岸阔叶林群系组(Formgroup Togay)
1.1.1 胡杨林群系(Form.Populus euphratica)
(1)胡杨-大叶白麻-胀果甘草群丛(Ass.P.euphratica-Poacynum hendersonii-Glycyrrhiza inflata)
(2)胡杨-芦苇-胀果甘草群丛(Ass.P.euphratica-Phragmites australis-G. inflata)
(3)胡杨-柽柳-胀果甘草群丛(Ass.P.euphratica-Tamarixsp.-G.inflata)
(4)胡杨-黑果枸杞-胀果甘草+黄甘草群丛(Ass.P.euphratica-Lycium ruthenicum-G.inflata+G.eurycarpa)
(5)胡杨-盐穗木-胀果甘草+花花柴群丛(Ass.P.euphratica-Halostachys caspica-G.inflata+Karelinia caspica)
(6)胡杨-铃铛刺-芨芨草-胀果甘草+光果甘草群丛(Ass.P.euphratica-Halimodendron halodendron-Achnatherum splendens-G.inflata+G.glabra)
(7)胡杨+灰杨-骆驼刺-光果甘草+胀果甘草群丛(Ass.P.euphratica+P. pruinosa-Alhagi sparsifolia-G.glabra+G.inflata)
1.1.2 灰杨林群系(Form.Populus pruinosa)
(1)灰杨+沙棘-光果甘草群丛(Ass.P.pruinosa+Hippophae rhamnoides-G. glabra)
(2)灰杨+胡杨-黑果枸杞-光果甘草群丛(Ass.P.pruinosa+P.euphratica-Lycium ruthenicum-G.glabra)
(3)灰杨-铃铛刺-光果甘草+胀果甘草群丛(Ass.P.pruinosa-Halimodendron halodendron-G.glabra+G.inflata)
1.1.3 白榆林群系(Form.Ulmus pumila)
(1)白榆+尖果沙枣-乌拉尔甘草群丛(Ass.U.pumila+Elaeagnus oxycarpa-G. uralensis)
(2)白榆-蒿-乌拉尔甘草群丛(Ass.U.pumila-Artemisiasp.-G.uralensis)
1.1.4 银灰杨林群系(Form.Populus canescens)
(1)银灰杨-野蔷薇-乌拉尔甘草群丛(Ass.P.canescens-Rosa sp.-G. uralensis)
(2)银灰杨+银白杨-油柴柳-乌拉尔甘草群丛(Ass.P.canescens+P.alba-Salix caspica-G.uralensis)
1.2 山地阔叶林群系组(Formgroup Broadleaf Forest)
1.2.1 山杨林群系(Form.Populus davidiana)
(1)山杨-虎榛子-东陵苔草-乌拉尔甘草群丛(Ass.P.davidiana-Ostryopsis davidiana-Carex tangiana-G.uralensis)
(2)山杨-甘肃山楂+达乌里胡枝子-乌拉尔甘草群丛(Ass.P.davidiana-Crataegus kansuensis+Lespedeza dahurica-G.uralensis)
1.3 山地针叶林群系组(Formgroup Needleleaf Forest)
1.3.1 油松林群系(Form.Pinus tabulaeformis)
(1)油松-胡枝子-乌拉尔甘草群丛(Ass.P.tabulaeformis-Lespedeza bicolor-G.uralensis)
(2)油松-白莲蒿-乌拉尔甘草群丛(Ass.P.tabulaeformis-Artimisia sacrorum-G.uralensis)
2 灌丛植被型(Scorub)
2.1 荒漠河岸灌丛群系组(Formgroup Togay)
2.1.1 黑果枸杞群系(Form.Lycium ruthenicum)
(1)黑果枸杞-大叶白麻-胀果甘草+黄甘草群丛(Ass.L.ruthenicum-Poacynum hendersonii-G.inflata+G.eurycarpa)
(2)黑果枸杞-花花柴-胀果甘草群丛(Ass.L.ruthenicum-Karelinia caspica-G.inflata)
(3)黑果枸杞-芦苇-胀果甘草群丛(Ass.L.ruthenicum-Phragmites australis-G.inflata)
2.1.2 多枝柽柳群系(Form.Tamarix ramosisima)
(1)多枝柽柳-胀果甘草群丛(Ass.T.ramosisima-G.inflata)
(2)多枝柽柳-铃铛刺-胀果甘草群丛(Ass.T.ramosisima-Halimodendron halodendron-G.inflata)
(3)多枝柽柳-假苇拂子茅-胀果甘草群丛(Ass.T.ramosisima-Calamagrostis pseudophragmites-G.inflata)
2.1.3 铃铛刺群系(Form.Halimodendron halodendron)
(1)铃铛刺-芨芨草-乌拉尔甘草群丛(Ass.H.halodendron-Achnatherum splendens-G.uralensis)
(2)铃铛刺-乌拉尔甘草+赖草群丛(Ass.H.halodendron-G.uralensis+Leymus secalinus)
2.2 荒漠盐柴类灌木群系组(Formgroup Halophyte Succulentoeremion Fsuticous)
2.2.1 盐穗木群系(Form.Halostachys caspica)
(1)盐穗木+黑果枸杞-乌拉尔甘草群丛(Ass.H.belangeriana+Lycium ruthenicum-G.uralensis)
(2)盐穗木-乌拉尔甘草-盐爪爪群丛(Ass.H.caspica-G.uralensis-Kalidium folitum)
(3)盐穗木-大叶白麻-胀果甘草(Ass.H.caspica-Poacynum hendersonii-G. inflata)
2.2.2 盐节木群系(Form.Halocnemum strobilaceum)
(1)盐节木-胀果甘草-盐生鸦葱群丛(Ass.H.strobilaceum-G.inflata-Scorzonera solsola)
(2)盐节木+刚毛柽柳-胀果甘草群丛(Ass.H.strobilaceum+Tamarix hispida-G.inflata)
(3)盐节木-芦苇-胀果甘草+花花柴群丛(Ass.H.strobilaceum-Phramites communis-G.inflata+Karelinia caspica)
2.3 荒漠蒿属小半灌木群系组(Formgroup Desert of Artemisia Microbushland)
2.3.1 白莲蒿群系(Form.Artemisia gmelisia)
(1)白莲蒿+乌拉尔甘草+长芒草群丛(Ass.A.gmelisia+G.uralensis+Stipa bungeana)
(2)白莲蒿-兴安胡枝子-乌拉尔甘草群丛(Ass.A.gmelisia-Lespedeza dahurica-G.uralensis)
(3)白莲蒿-乌拉尔甘草群丛(Ass.A.gmelisia-G.uralensis)
2.3.2 油蒿群系(Form.Artemisia ordosica)
(1)乌拉尔甘草+黑沙蒿-骆驼蒿+沙米群丛(Ass.G.uralensis+A.ordosica-Feganum harmala+Agriophyllum squarrosum)
(2)油蒿+乌拉尔甘草—无芒隐子草+刺沙蓬群丛(Ass.A.ordosica+G.uralensis-Cleistogenes songoarica+Salsola ruthenica)
(3)黑沙蒿+冷蒿+乌拉尔甘草-沙生针茅+沙兰刺头群丛(Ass.A.ordosica+A. frigida+G.uralensis-Stipaglareosa+Echinops gmelinii)
2.4 荒漠草原化灌木群系组(Formgroup Desert Steppe Fruticous)
2.4.1 锦鸡儿群系(Form.Caraganasp.)
(1)柠条锦鸡儿-乌拉尔甘草群丛(Ass.Caragana korshinskii-G.uralensis)
(2)藏锦鸡儿-乌拉尔甘草-戈壁针茅群丛(Ass.C.tibetica-G.uralensis-Stipa tianschanica subsp.gobica)
3 草甸植被型(Meadow)(www.xing528.com)
3.1 亚高山及山地草甸群系组(Formgroup Subalpine and Alpine Meadow)
3.1.1 根茎禾草群系(Form.rhizome agrosto group)
(1)黄菅草-白草-乌拉尔甘草群丛(Ass.Themeda triandra var.japonica-Bothriochloa ischaemum-G.uralensis)
(2)野古草-乌拉尔甘草群丛(Ass.Arundinella hirta-G.uralensis)
(3)无芒燕麦-甘草群丛(Ass.Avena fatua var.glabrata-G.uralensis)
3.2 低地、河漫滩真草甸群系组(Formgroup Lowland Flood of True Meadow)
3.2.1 根茎禾草群系(Form.rhizome agrosto group)
(1)假苇拂子茅-乌拉尔甘草群丛(Ass.Calamagrostis pseudophragmites-G. uralensis)
(2)小糠草-乌拉尔甘草群丛(Ass.Agrotis alba-G.uralensis)
(3)偃麦草-乌拉尔甘草群丛(Ass.Elytrigia repens-G.uralensis)
3.3 低地、河漫滩盐化草甸群系组(Formgroup Lowland Flood Meadow of Salinization)
3.3.1 芨芨草群系(Form.Achnatherum splendens)
(1)芨芨草-赖草+乌拉尔甘草群丛(Ass.A.splendens-Leymus secalinus+G. uralensis)
(2)芨芨草-拂子茅-乌拉尔甘草群丛(Ass.A.splendens-Calamagrostis epigeios-G.uralensis)
(3)芨芨草-乌拉尔甘草群丛(Ass.A.splendens-G.uralensis)
(4)芨芨草-乌拉尔甘草-碱茅群丛(Ass.A.splendens-G.uralensis-Puccinellia distans)
(5)乌拉尔甘草-芨芨草-芦苇群丛(Ass.G.uralensis-A.splendens-Phragmites communis)
3.3.2 芦苇群系(Form.Phragmites australis)
(1)芦苇-假苇拂子茅-乌拉尔甘草群丛(Ass.P.australis-Calamagrostis pseudophragmites-G.uralensis)
(2)芦苇-大叶白麻-乌拉尔甘草群丛(Ass.P.communis-Poacynum hendersonii-G.uralensis)
(3)芦苇-大叶白麻-光果甘草群丛(Ass.P.communis-Poacynum hendersonii-G.glabra)
(4)芦苇-胀果甘草群丛(Ass.P.australis-G.inflata)
3.3.3 胀果甘草群系(Form.Glycyrrhiza inflata)
(1)胀果甘草-花花柴群丛(Ass.G.inflata-Karelinia caspica)
(2)胀果甘草-盐地风毛菊群丛(Ass.G.inflata-Saussurea salsa)
(3)胀果甘草-小獐茅群丛(Ass.G.inflata-Aeluropus pungens)
(4)胀果甘草-海乳草+水麦冬群系(Ass.G.inflata-Glaux maritima+Triglocbin palustre)
3.3.4 杂类草群系(Form.weeds)
(1)大叶白麻-胀果甘草群丛(Ass.Poacynum hendersonii-G.inflata)
(2)大叶白麻-乌拉尔甘草+光果甘草群丛(Ass.P.hendersonii-G.uralensis+G. glabra)
(3)骆驼刺-胀果甘草群丛(Ass.Alhagi sparsifolia-G.inflata)
(4)花花柴-胀果甘草-狗牙根群丛(Ass.Karelinia caspica-G.inflata-Cynodon dactylon)
4 平原洲及灌区植被型(Planit)
4.1 绿洲阔叶林群系组(Formgroup Oasis Broadleaf Forest)
4.1.1 杨树林群系(Form.Populus sp.)
(1)钻天杨-假苇拂子茅-乌拉尔甘草群丛(Ass.P.nigra var.italica-Calamagrostis pseudophragmites-G.uralensis)
(2)银白杨-光果甘草-黄花草木樨群丛(Ass.P.alba-G.glabra-Melilotus officinalis)
4.1.2 白腊树林群系(Form.Fraxinus rhynchophylla)
(1)白腊-光果甘草+乌拉尔甘草-苦马豆群丛(Ass.Fraxinus rhynchophylla-G. glabra+G.uralensis-Trifolium repens)
(2)白腊+白榆-光果甘草群丛(Ass.F.rhynchophylla+Ulmus pumila-G. glabra)
4.1.3 苹果树群系(Form.Malus pumila)
(1)苹果-假苇拂子茅-光果甘草群丛(Malus pumila-Calamagrostis pseudophragmites-G.glabra)
(2)苹果-假苇拂子茅+光果甘草群丛(Ass.M.pumila-Calamagrostis pseudophragmites+G.glabra)
4.1.4 葡萄群系(Form.Vitis vinifera)
(1)葡萄-苦豆子+蒙山莴苣-粗毛甘草群丛(Ass.V.vinifera-Sophora alopecuroides+Lactuca tatarica-G.aspera)
(2)葡萄-假苇拂子茅-粗毛甘草群丛(Ass.V.vinifera-Calamagrostis pseudophragmites-G.aspera)
4.1.5 沙枣群系(Form.Elaeagnus oxycarpa)
(1)尖果沙枣-苦马豆+胀果甘草群丛(Ass.E.oxycarpa-Swainsonia salsula+G. inflata)
(2)尖果沙枣-芦苇-黄甘草群丛(Ass.E.oxycarpa-Phragmites australis-G. eurycarpa)
(3)尖果沙枣-乌拉尔甘草-沙生旋覆花(Ass.E.oxycarpa-G.uralensis-Inula salsoloides)
4.2 杂类草群系组(Formgroup weeds)
4.2.1 粗毛甘草群系(Form.Glycyrrhiza aspera)
(1)粗毛甘草-角果藜群丛(Ass.G.aspera-Ceratocarpus arenarius)
(2)粗毛甘草-灰绿藜群丛(Ass.G.aspera-Chenopodium glaucum)
(3)粗毛甘草-小旋花群丛(Ass.G.aspera-Convolvulus arvensis)
4.2.2 黄花草木樨群系(Form.Melilotus officinalis)
(1)黄花草木樨+乌拉尔甘草-白花三叶草群丛(Ass.M.officinalis+G.uralensis-Triffolium repens)
(2)黄花草木樨+光果甘草-狗尾草群丛(Ass.M.officinalis+G.glabra-Setaria viridis)
4.2.3 苦苣菜群系(Form.Sonchus oleraceus)
(1)苦苣莱+胀果甘草-道爪草群丛(Ass.S.oleraceus+G.inflata-Dodartia orientalis)
(2)苦苣莱+光果甘草-蒲公英群丛(Ass.S.oleraceus+G.glabra-Taraxacum mongolicum)
4.3 农作物群系组(Formgroup Crops)
4.3.1 蔬菜群系(Form.vagetable)
4.3.2 禾谷群系(Form.grain)
4.3.3 棉花群系(Form.cotton)
李学禹 陆源芬 阎平
国家自然科学基金项目:3870031
发表于《石河子农学院学报》,1991,18(3):1-6
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