王會(huì)儒,陳國(guó)鵬,2*,王 飛,曹秀文,郭 星
(1.甘肅省白龍江林業(yè)管理局 林業(yè)科學(xué)研究所,甘肅 武都746010;2.甘肅農(nóng)業(yè)大學(xué) 林學(xué)院,甘肅 蘭州730070)
干旱河谷(dry valley),是指在高山峽谷區(qū)一些深切河谷的下部出現(xiàn)的四周被相對(duì)濕潤(rùn)環(huán)境所包圍的較干旱、溫度較高、以灌叢和草叢植被為主的河谷生態(tài)系統(tǒng)或河谷景觀。它與周邊地區(qū)濕潤(rùn)、半濕潤(rùn)等景觀不相協(xié)調(diào),是青藏高原南、東、北邊緣的高山峽谷地帶植被退化嚴(yán)重的獨(dú)特的生態(tài)景觀[1]。干旱河谷植被稀疏且退化嚴(yán)重,是典型的生態(tài)脆弱帶,且具有荒漠化或石漠化的趨勢(shì)[2]。由于干旱河谷主要分布于長(zhǎng)江上游主要支流的中上游,生態(tài)地位十分顯著,其生態(tài)安全是保障長(zhǎng)江上游生態(tài)屏障的必須;同時(shí),該區(qū)域也是我國(guó)西部經(jīng)濟(jì)建設(shè)的支撐點(diǎn),其生態(tài)環(huán)境的惡化不僅威脅著區(qū)域安全,甚至?xí){長(zhǎng)江中下游地區(qū)的穩(wěn)定和發(fā)展[3]。
多年來(lái),干旱河谷脆弱環(huán)境一直是林學(xué)家、生態(tài)學(xué)家和各級(jí)政府關(guān)注的焦點(diǎn)。針對(duì)我國(guó)干旱河谷生態(tài)系統(tǒng)特殊性的植被恢復(fù)與生態(tài)系統(tǒng)管理研究已開(kāi)展多年。研究?jī)?nèi)容從最初的地質(zhì)、地貌形成和氣候特性領(lǐng)域揭示干旱河谷成因逐漸擴(kuò)展到了生物多樣性及其功能適應(yīng)機(jī)制的方向、退化生態(tài)系統(tǒng)的恢復(fù)和重建。國(guó)家近幾十年來(lái)陸續(xù)實(shí)施了長(zhǎng)江上游水土流失重點(diǎn)防治、長(zhǎng)江中上游防護(hù)林體系建設(shè)、天然林保護(hù)、退耕還林等重大生態(tài)工程,相關(guān)研究和工程的實(shí)施使干旱河谷生態(tài)系統(tǒng)惡化的趨勢(shì)得以控制,在恢復(fù)模式、生態(tài)系統(tǒng)管理、人工植被的效益評(píng)價(jià)等方面取得了較多突破,已篩選出一批樹(shù)(草)種,有效地恢復(fù)了部分地區(qū)的植被覆蓋[4]。由于長(zhǎng)期重視恢復(fù)實(shí)踐而忽視基礎(chǔ)數(shù)據(jù)積累和基礎(chǔ)理論總結(jié),使植被恢復(fù)經(jīng)驗(yàn)很難在大區(qū)域推廣且植被恢復(fù)后的長(zhǎng)期效應(yīng)不確定,干旱河谷植被恢復(fù)難題尚未攻破[5]。鑒于此,從植物個(gè)體、種群、群落和生態(tài)系統(tǒng)到景觀尺度系統(tǒng)總結(jié)了干旱河谷植物適應(yīng)機(jī)制和植被恢復(fù)重建取得的基礎(chǔ)成果,提出了未來(lái)應(yīng)予重視的研究方向,以期為干旱河谷植被恢復(fù)重建的研究與實(shí)踐提供參考。
干旱河谷主要分布在橫斷山脈的怒江、瀾滄江、金沙江和雅礱江的中、下游,大渡河和元江的中游,岷江和嘉陵江上游,北起白龍江上游谷地(32°21′N),南至元江谷地(22°33′N),東起金沙江下游及大渡河谷地,西至藏東的昌都、八宿一線,總長(zhǎng)4 105 km,總面積約11 230km2[6]。干旱河谷所處的山脈河流多作南北走向,平行相間排列,與控制這一地區(qū)的西南季風(fēng)形成較大的交角,水汽多在迎風(fēng)坡凝結(jié)降落,造成河谷降水偏低;此外由于河谷地形閉塞,太陽(yáng)輻射強(qiáng)烈,散熱不宜,加之焚風(fēng)效應(yīng),呈現(xiàn)出溫度高、蒸發(fā)強(qiáng)、干旱重的氣候特點(diǎn)。由于分布區(qū)域地理跨度大,干旱河谷氣候多樣。南部河谷年均溫20℃以上,北部河谷年均溫6~13℃,氣溫年較差小,日較差大,春季氣溫回升快,秋季下降迅速。多數(shù)地方降雨量介于500~800mm,集中在5-10月,干濕季明顯;蒸發(fā)強(qiáng)烈,年蒸發(fā)量是降雨量的1.5~3.6倍[7]。根據(jù)最冷月平均氣溫、日均溫≥10℃的持續(xù)天數(shù)、年積溫等熱量條件,又可將干旱河谷劃分為干熱河谷、干暖河谷、干溫河谷和干涼河谷[6]。
干旱河谷所處橫斷山區(qū)地質(zhì)構(gòu)造活躍,地震、山體滑坡、泥石流等自然災(zāi)害多發(fā),進(jìn)而加劇脆弱生態(tài)生態(tài)系統(tǒng)的易變性。災(zāi)害不僅損毀了河谷稀疏植被和稀薄土壤,導(dǎo)致異質(zhì)的景觀單元[8];還會(huì)影響植被的恢復(fù)潛力,如災(zāi)害堆積物破壞土壤種子庫(kù),阻斷了植被的自然更新[9]。干旱河谷生態(tài)系統(tǒng)受損程度與自然災(zāi)害發(fā)生的強(qiáng)度、頻度、過(guò)程和持續(xù)時(shí)間等息息相關(guān)。干旱河谷的巖土類型多樣、土被不連續(xù)、坡面土層淺薄、巖石裸露率高等造就了干旱河谷土壤強(qiáng)烈的水分、養(yǎng)分空間分異,高度異質(zhì)的土壤環(huán)境影響著植物的生長(zhǎng)、生物量積累和資源利用等生理生態(tài)過(guò)程[10],并決定著天然植被的分布[11]。宋成軍[12]等就發(fā)現(xiàn)低水、低磷和高氮抑制小馬鞍羊蹄甲(Bauhinia faberi)的存活與生長(zhǎng),薛沛沛[13]等指出當(dāng)土壤含水率為田間持水量的70%~85%時(shí)才能很好地促進(jìn)臺(tái)灣青棗(Zizyphus mauritiana)生長(zhǎng)。
土壤滲透性決定土壤吸收外界供水(降水)的能力,奠定土地對(duì)植物可供水的容量基礎(chǔ),是支持干旱河谷植物成活和生長(zhǎng)的土壤水分性質(zhì)中最關(guān)鍵因素[14],其與巖土性質(zhì)密切相關(guān),泥巖坡地土體黏重板結(jié),入滲能力弱;片巖坡地、礫石巖坡地、砂礫巖坡地等石質(zhì)坡地土體裂縫發(fā)育,入滲能力強(qiáng)[15],恢復(fù)植被將有效地改善土壤的滲透性能[16-17]。此外,干旱河谷土壤粒級(jí)小,斥水性強(qiáng)度與分布比例高,導(dǎo)致嚴(yán)重水土流失[18]、土壤砂礫化[19],進(jìn)而使土壤更加貧瘠[20]。
干旱河谷獨(dú)特的環(huán)境條件造就了獨(dú)特的植被,其類型、種類組成、群落及其動(dòng)態(tài)復(fù)雜多樣,且多為非地帶性植被。現(xiàn)存植被主要為灌叢、草叢,群落結(jié)構(gòu)單一,植物種類以陽(yáng)性為主,呈多毛、具刺、葉小、質(zhì)厚、低矮或匍匐生長(zhǎng)等典型旱生特征[21-22]。從植物區(qū)系看,南部干熱河谷以喜干熱的熱帶成分為主體[23-24],北部白龍江干溫河谷植物具有明顯的溫帶屬性,并在起源上與熱帶、亞熱帶植物有一定淵源[22]。
植物對(duì)干旱河谷適應(yīng)性的報(bào)道較多,以適應(yīng)干旱脅迫的研究為主,涉及形態(tài)解剖、生理生化過(guò)程、水分利用方式和途徑等。以落葉、植株矮化、枝葉刺化、葉片膜質(zhì)或鱗片化、葉質(zhì)硬化和減小葉面積為主要形態(tài)適應(yīng)途徑[21],在生理生化過(guò)程中,干旱河谷植物通過(guò)有效的吸水能力和完善的保水機(jī)制來(lái)提高水分利用效率和進(jìn)行有利的光保護(hù)機(jī)制適應(yīng)[25-28];如段愛(ài)國(guó)[29]等就發(fā)現(xiàn)干旱河谷植物的表觀光合量子效率均低于自然界最大量子效率的理論值。在應(yīng)對(duì)干旱脅迫過(guò)程中,植物適應(yīng)機(jī)制表現(xiàn)為多態(tài)性,如K+和可溶性糖在干旱的前后期貢獻(xiàn)較大[28],游離脯氨酸、游離氨基酸在干旱的中期起滲透調(diào)節(jié)作用[30];光合和蒸騰速率的物種差異較大[31],有增強(qiáng)型[32]、減弱性及穩(wěn)定型等形式[33],優(yōu)勢(shì)植物體內(nèi)的谷胱甘肽轉(zhuǎn)移酶和谷胱甘肽過(guò)氧化物酶在干熱季能特異性表達(dá)[34]。
海拔跨度大,坡向、坡位效應(yīng)明顯,是顯著影響干旱河谷植物表型的主要環(huán)境因子。研究發(fā)現(xiàn),葉面積、葉厚度、柵欄組織和海綿組織的厚度、氣孔器面積、表皮細(xì)胞面積以及表皮毛密度隨海拔升高呈增加趨勢(shì),而角質(zhì)膜厚度、表皮細(xì)胞密度和表皮毛長(zhǎng)度的變化則 相 反[35];岷江 柏 木 (Cupressus chengiana)會(huì)通過(guò)降低光合速率、水分利用效率和生理內(nèi)含物適應(yīng)海拔升高[36]。坡位對(duì)植物的生理過(guò)程影響顯著,段愛(ài)國(guó)等發(fā)現(xiàn)低坡位有助于植物維持相對(duì)高的凈光合速率、促進(jìn)蒸騰,且低坡位的光合增益效應(yīng)在干熱季節(jié)更明顯[37]。
干旱河谷植物種群生態(tài)學(xué)的研究主要體現(xiàn)在種群繁殖策略、種群結(jié)構(gòu)與動(dòng)態(tài)、種群競(jìng)爭(zhēng)、生物量構(gòu)成與分配格局等方面,目前取得的成果有:多苞薔薇(Rosa multibracteata)能通過(guò)有性結(jié)實(shí)與無(wú)性萌蘗生殖方式的調(diào)整來(lái)平衡資源的分配利用以適應(yīng)多變環(huán)境,在適應(yīng)過(guò)程中還能表現(xiàn)出良好的種群持續(xù)性[38];砂生槐(Sophora moorcroftiana)對(duì)雅魯藏布江干旱河谷適應(yīng)過(guò)程中表現(xiàn)出種群個(gè)體集中在中等大小范圍為主,種群的發(fā)展會(huì)經(jīng)過(guò)少數(shù)個(gè)體集群性定居、在原有聚集尺度范圍繁殖到種群改良局部環(huán)境,密度增大,種內(nèi)競(jìng)爭(zhēng)加劇,最終種群向更大尺度擴(kuò)散和壯大[39]。干旱河谷種群的分布除受自然因素作用外,還受動(dòng)物采食影響,如錐花小檗(Berberis prattii)種群一般集中分布在遠(yuǎn)離牧道的區(qū)域[40]。
在水分虧缺,養(yǎng)分貧瘠的干旱河谷,樹(shù)木為了保持高的組織水勢(shì)和正常生長(zhǎng),將采取發(fā)達(dá)根系的策略來(lái)吸收更廣范圍的土壤水分和養(yǎng)分,較多的研究采用生物量與生產(chǎn)力的變化來(lái)評(píng)價(jià)種群的可適性[41],如赤桉(Eucalyptus camaldulensis)和新銀合歡樹(shù)(Leucaena leucocephala)將更多的生物資源投入到根系構(gòu)建中,其分配到干的生物量比例低于我國(guó)大多數(shù)人工林[42]。干旱河谷深根系的植物根幅大[43]、總根長(zhǎng)和根系生物量高[44]、資源利用持續(xù)時(shí)間長(zhǎng)[45],而垂直分布淺的植物表現(xiàn)為發(fā)達(dá)細(xì)根的適應(yīng)策略[46]。
干旱河谷植被的頂級(jí)群落尚未統(tǒng)一,有學(xué)者認(rèn)為人類活動(dòng)是導(dǎo)致干旱河谷植被群落退化的主要原因[47];而 Moseley和唐亞通過(guò)比對(duì)歷史照片,發(fā)現(xiàn)過(guò)去150a間干旱河谷植被范圍并沒(méi)有顯著的變化[48]。就干旱河谷現(xiàn)存植被而言,植被類型組合及群落演替方向均具有較大的波動(dòng)性和不確定性,次生演替現(xiàn)象明顯[49-50]、群落層次結(jié)構(gòu)單一、群落外貌隨干濕季交替變化明顯[51],在谷坡上的分布呈斷續(xù)狀[23],并認(rèn)為水分主導(dǎo)干旱河谷植物群落分布,影響群落內(nèi)不同層次物種數(shù)量結(jié)構(gòu)和空間格局[52],而海拔和坡向、坡位等通過(guò)影響著土壤的養(yǎng)分狀況進(jìn)而影響群落的多樣性和穩(wěn)定性[53-57]。
種群對(duì)不同資源空間的利用能力和適應(yīng)性不盡相同[58],引起的種群分布范圍和均勻程度的分化,將決定群落中優(yōu)勢(shì)種的構(gòu)成。物種功能性狀的組配和體內(nèi)元素的耦聯(lián)在群落構(gòu)建中起著重要意義,主要的研究涉及干旱河谷植物功能性狀空間差異、元素含量在群落構(gòu)建中的作用等[59]。如Fe在干旱河谷群落構(gòu)建上作用重要,P在植物群落生產(chǎn)力和群落構(gòu)建中作用顯著[60];而在金沙江干熱河谷,葉片N、P含量只有全國(guó)平均值的1/2[61],K 接近于全國(guó)均值10.3mg·g-1,Ca含量高于全國(guó)平均水平[62]。生態(tài)系統(tǒng)恢復(fù)會(huì)降低植物葉片N、P含量以及P/K,改變計(jì)量關(guān)系,進(jìn)一步引起群落物種優(yōu)勢(shì)度的變化[63]。
在群落自然演替的過(guò)程中,土壤種子庫(kù)是基礎(chǔ),植物株型是影響種子庫(kù)的主要因素[64-66],而泥石流干擾將嚴(yán)重破壞土壤種子庫(kù)[9]。種子的萌發(fā)需要濕潤(rùn)的環(huán)境保障[67];在應(yīng)對(duì)惡劣環(huán)境是,種子能通過(guò)休眠方式使其在萌發(fā)的時(shí)間和空間上分散,形成大容量的種子庫(kù),確保部分種子的萌發(fā)和存活[68-69]。盡管干旱河谷灌叢具有依靠土壤種子庫(kù)實(shí)現(xiàn)自我更新的潛力,但由于物種組成的Sorensen相似性較低,現(xiàn)存灌叢植被一旦遭到破壞后,單依靠土壤種子庫(kù)自然恢復(fù)到現(xiàn)存植被比較困難[70]。在人工恢復(fù)植被的過(guò)程中,進(jìn)行封育禁牧可以提高了土壤種子庫(kù)密度[68];小容器播種育苗、保水劑和腐殖質(zhì)等土壤改良措施可以提高種子的出苗率,但不能提高幼苗最終的存活率和生長(zhǎng)速率[67]。
生態(tài)條件的特殊性決定了干旱河谷群落/生態(tài)系統(tǒng)的較低生物量積累,在岷江干旱河谷的對(duì)節(jié)刺(Sageretia pycnophylla)灌叢地上生物量達(dá)1.16t·hm-2,是幾種灌叢中最高的,卻遠(yuǎn)低于地帶性植被群落生物量[57]。植被恢復(fù)過(guò)程中引進(jìn)外來(lái)樹(shù)種的生產(chǎn)力和生物量積累明顯高于本地灌叢[71],但其穩(wěn)定性尚需持續(xù)關(guān)注。雖然凋落物動(dòng)態(tài)分解在生態(tài)系統(tǒng)物質(zhì)循環(huán)和功能發(fā)揮方面作用突出,但干旱河谷的相關(guān)研究鮮見(jiàn)報(bào)道。
生態(tài)系統(tǒng)物質(zhì)循環(huán)和能量流動(dòng)過(guò)程的研究是揭示系統(tǒng)功能和制定干旱河谷綜合整治方案的關(guān)鍵,對(duì)干旱河谷植物的水能利用、光能利用等資源利用的研究?jī)H停留在個(gè)體尺度[27,29],尚不足以推演到群落和生態(tài)系統(tǒng)尺度。
從景觀生態(tài)學(xué)的視角開(kāi)展干旱河谷的研究主要集中于岷江上游的土地利用變化、微尺度空間格局、干擾事件等領(lǐng)域。楊兆平[72]等發(fā)現(xiàn)干旱河谷的整體形狀簡(jiǎn)單,土地利用類型分布呈垂直帶型[73],灌木林地是景觀基質(zhì),占60%以上面積[32],植被景觀由川甘亞菊(Ajania potaninii)、刺蓬(Herbal euphorbiae)、瓦松(Orostachys fimbriatus)等10多種斑塊控制,植物分布呈現(xiàn)較高空間聚集性[74]。近年來(lái),干旱河谷景觀破碎化程度和異質(zhì)性程度持續(xù)增加,斑塊密度、多樣性指數(shù)、蔓延度指數(shù)、邊界密度和分維數(shù)等在1995年前后變化各異[75],人口增加和國(guó)家政策驅(qū)動(dòng)岷江干旱河谷景觀變化[32],地震干擾影響干旱河谷景觀的穩(wěn)定[76]。
多年來(lái)干旱河谷的植被恢復(fù)一直得到各界的重視,通過(guò)選育鄉(xiāng)土樹(shù)種、引種樹(shù)種造林和封禁修復(fù)等多項(xiàng)措施,并在各地建立了許多快速恢復(fù)植被的模式,干旱河谷植被退化的勢(shì)頭在一些地方得到了初步遏制,植被覆蓋度和土壤肥力等有所提高。但是,由于對(duì)干旱河谷植被演替規(guī)律認(rèn)識(shí)不足、植被恢復(fù)多以營(yíng)造人工林為主等,部分早期被認(rèn)為最有效的治理模式目前也暴露出了許多問(wèn)題,取得的初步成效能否持續(xù)還有待進(jìn)一步檢驗(yàn)[5],干旱河谷植被恢復(fù)的任務(wù)依然艱難,未來(lái)的研究中還需要關(guān)注以下幾個(gè)問(wèn)題。
1)干旱河谷植被演替及退化機(jī)制。植物群落演替規(guī)律是制定干旱河谷植被恢復(fù)宏觀決策和具體措施的理論基礎(chǔ),自然恢復(fù)是干旱河谷生態(tài)系統(tǒng)恢復(fù)與重建最重要和切實(shí)可行的途徑。目前,關(guān)于干旱河谷群落演替規(guī)律的報(bào)道還表現(xiàn)在對(duì)各演替階段現(xiàn)象的描述,鮮有報(bào)道涉及群落演替動(dòng)力和機(jī)理,涉及的群落類型亦不全面。未來(lái)應(yīng)該在不同氣候帶、不同巖性構(gòu)成和不同利用方式和強(qiáng)度的地方,結(jié)合生態(tài)系統(tǒng)過(guò)程的研究探索群落演替的規(guī)律和機(jī)制,揭示群落穩(wěn)定性維持的機(jī)理和限制退化生態(tài)系統(tǒng)恢復(fù)的主導(dǎo)因素并尋求解決的途徑。同時(shí),干旱河谷植物如何適應(yīng)高度異質(zhì)性生境的研究仍需高度重視,如一些克隆植物的覓養(yǎng)行為、不同資源利用型植物之間的組合關(guān)系等。
2)從植物功能屬性為出發(fā)點(diǎn)認(rèn)識(shí)干旱河谷植被生態(tài)學(xué)過(guò)程,注重植物適應(yīng)機(jī)制在葉片-個(gè)體-種群-群落/生態(tài)系統(tǒng)-景觀不同尺度間的推演。干旱河谷微生境對(duì)功能屬性的影響顯著,從功能屬性組裝配置了解植物在不同尺度的適應(yīng)機(jī)制。在方法學(xué)來(lái)看,干旱河谷生態(tài)系統(tǒng)根系的研究一直是難點(diǎn)之一。生境高度異質(zhì)性,土壤質(zhì)地和基質(zhì)的變化極為復(fù)雜,且土壤養(yǎng)分和水分分布的極不均勻,植物的根冠分配隨生境的不同而有較大分化,植物的根系多生長(zhǎng)在巖隙間,挖掘和測(cè)定相當(dāng)困難。所以,未來(lái)需要繼續(xù)探索同位素示蹤或超聲探測(cè)來(lái)研究植物的根系功能,借鑒醫(yī)學(xué)X-光、B-超等技術(shù)進(jìn)行葉片功能和樹(shù)皮功能屬性研究。另外,以生態(tài)化學(xué)計(jì)量特征為出發(fā)點(diǎn)闡明干旱河谷生態(tài)系統(tǒng)土壤盈虧與養(yǎng)分限制,揭示其內(nèi)穩(wěn)性機(jī)制。
3)頻發(fā)的地質(zhì)災(zāi)害對(duì)植被及其生態(tài)效益的影響。如地質(zhì)災(zāi)害的對(duì)植被生態(tài)系統(tǒng)的影響,植物受災(zāi)后的自我修復(fù)機(jī)制、災(zāi)害跡地殘存植被的保育技術(shù),植被的水文過(guò)程與功能、植被的減災(zāi)效應(yīng)等內(nèi)容也應(yīng)在今后干旱河谷植被生態(tài)學(xué)的研究中得到足夠的重視。
4)干旱河谷適生經(jīng)濟(jì)植物的開(kāi)發(fā)利用。干旱河谷特殊的環(huán)境條件塑造了許多植物特殊的經(jīng)濟(jì)品相,如干旱河谷是引種油橄欖(Olea europaea)的唯一適生區(qū)。在植被恢復(fù)過(guò)程中探索經(jīng)濟(jì)植物的高效栽培措施、注重特殊功能器官的開(kāi)發(fā)利用研究,促進(jìn)生態(tài)建設(shè)和經(jīng)濟(jì)建設(shè)共贏,鞏固生態(tài)建設(shè)成果。
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