段后浪,趙 安①,姚 忠
(1.江西師范大學(xué)鄱陽(yáng)湖濕地與流域研究教育部重點(diǎn)實(shí)驗(yàn)室,江西 南昌 330022;2.江西師范大學(xué)地理與環(huán)境學(xué)院,江西 南昌 330022;3.江西省科學(xué)院,江西 南昌 330096)
鄱陽(yáng)湖濕地典型草洲主要植物種群在5個(gè)資源環(huán)境梯度上的生態(tài)位特征
段后浪1,2,趙 安1,2①,姚 忠3
(1.江西師范大學(xué)鄱陽(yáng)湖濕地與流域研究教育部重點(diǎn)實(shí)驗(yàn)室,江西 南昌 330022;2.江西師范大學(xué)地理與環(huán)境學(xué)院,江西 南昌 330022;3.江西省科學(xué)院,江西 南昌 330096)
鄱陽(yáng)湖典型濕地;植物種群;生態(tài)位寬度;生態(tài)位重疊
“生態(tài)位理論”是解析植物群落物種組成變化主要機(jī)制的理論之一,研究植物種群生態(tài)位對(duì)于理解植物群落結(jié)構(gòu)和功能、群落內(nèi)物種間關(guān)系、生物多樣性、群落動(dòng)態(tài)演替和種群進(jìn)化等方面有著重要意義[1]。目前,對(duì)于植物種群生態(tài)位的研究主要包括生態(tài)位寬度和生態(tài)位重疊,大都采用Levins生態(tài)位寬度指數(shù)及Pianka生態(tài)位重疊指數(shù),模型相對(duì)簡(jiǎn)單,揭示的生態(tài)學(xué)意義明顯[2-4]。
恒湖農(nóng)場(chǎng)茶葉港草洲位于鄱陽(yáng)湖畔,屬鄱陽(yáng)湖沖積平原。洲灘植被具有重要的生態(tài)功能,不僅在生物多樣性保育、生態(tài)系統(tǒng)穩(wěn)定性的維持等方面發(fā)揮重要作用,而且還為成千上萬(wàn)只越冬候鳥(niǎo)提供棲息地和食物(植物莖、葉等),研究鄱陽(yáng)湖典型草洲植物種群的生態(tài)位特征對(duì)植被資源保護(hù)、可持續(xù)利用和植被恢復(fù)重建等具有重要意義。目前,針對(duì)鄱陽(yáng)湖區(qū)域草洲植物種群生態(tài)位的研究鮮有報(bào)道,僅陳明華等[5]于2011年在鄱陽(yáng)湖北部星子縣蓼花洲、中部吳城和南部的南磯山國(guó)家級(jí)自然保護(hù)區(qū)做了該方面的分析探索,該文基于植物特征(高度、蓋度、生物量)進(jìn)行了生態(tài)位研究,但沒(méi)有涉及環(huán)境梯度上的生態(tài)位特征研究,很難在不同植物種群生態(tài)位差異機(jī)制上給出合理解釋。筆者基于鄱陽(yáng)湖濕地恒湖農(nóng)場(chǎng)茶葉港草洲19個(gè)樣地95個(gè)樣方植物與5個(gè)資源環(huán)境梯度(高程、土壤pH值、電導(dǎo)率、含水量和有機(jī)質(zhì)含量)數(shù)據(jù),分析不同資源環(huán)境因子梯度上研究區(qū)主要植物種群生態(tài)位寬度、生態(tài)位重疊情況以及物種生態(tài)位寬度與重要值及其變異系數(shù)間的相關(guān)程度,探索主要物種對(duì)不同環(huán)境因子變化的適應(yīng)性,揭示各物種在群落中的功能和地位,為鄱陽(yáng)湖濕地草洲植物物種多樣性保護(hù)及植被恢復(fù)提供科學(xué)依據(jù)。
1.1 研究區(qū)概況
1.2 研究方法
1.2.1 數(shù)據(jù)采集
正式調(diào)查之前在恒湖農(nóng)場(chǎng)茶葉港草洲進(jìn)行相關(guān)預(yù)試驗(yàn),確定能代表植物群落多樣性和分布特征的具體范圍。于2015年10月底,按大致垂直于湖岸線的地形梯度平行設(shè)置4條樣帶,樣帶間距為80 m,每條樣帶等間距60 m選取5個(gè)5 m×5 m的樣地,按X型在樣地四角及中心取5個(gè)1 m×1 m的樣方,數(shù)據(jù)采集以樣方為單位。(1)植被數(shù)據(jù)及地形:記錄每個(gè)樣方的植物種類(lèi),用米尺測(cè)定樣方中物種株高,取最高值為植物高度,以植物地上部分的垂直投影面積與樣方面積比值為植物蓋度,用GPS進(jìn)行定位并記錄立地微環(huán)境。(2)部分土壤因子現(xiàn)場(chǎng)測(cè)定:每個(gè)樣方中心用土壤三參數(shù)傳感器GS3(美國(guó)DECAGON公司生產(chǎn))測(cè)定0~15 cm土層[6]土壤含水量及電導(dǎo)率。(3)土樣采集及實(shí)驗(yàn)室處理:用取土環(huán)刀采集每個(gè)樣方中心0~15 cm土層土樣約100 g,將同一樣地5份土樣充分混合為1份。將樣品自然風(fēng)干后,撿去植物殘根及碎石塊等雜物,磨碎后過(guò)149 μm孔徑篩,參考賈建麗等[7]實(shí)驗(yàn)室土壤指標(biāo)測(cè)定方法:有機(jī)質(zhì)含量采用重鉻酸鉀氧化外加熱法,pH值采用電位法(水土體積比為5∶1)。其中,第1條樣帶中的3號(hào)樣地所在位置因出現(xiàn)一片較大人工水塘,沒(méi)有采樣。
1.2.2 數(shù)據(jù)處理
(1)重要值及其變異系數(shù)計(jì)算
植物物種重要值是反映物種在植物群落中的作用和地位的綜合數(shù)量指標(biāo)。目前,有關(guān)重要值的計(jì)算方法很多且不同植物層次(喬木、灌木、草本等)也大都不一,以19個(gè)樣地(5 m×5 m)為基礎(chǔ),樣地中每種草本植物的重要值計(jì)算公式[8]為
P=(cr+hr+fr)/3。
(1)
式(1)中,P為樣地中每種植物重要值;cr為相對(duì)蓋度,指每個(gè)樣地中每種植物蓋度占該樣地中所有植物總蓋度的比例,樣地中每種植物蓋度由該樣地5個(gè)樣方該種植物蓋度取平均值得到;hr為相對(duì)高度,計(jì)算方法與相對(duì)蓋度計(jì)算方法類(lèi)似;fr為相對(duì)頻度,指樣地中每種植物在5個(gè)樣方中的出現(xiàn)頻度占所有物種總頻度的比例。
變異系數(shù)是衡量各觀測(cè)值變異程度或離散程度的另一個(gè)統(tǒng)計(jì)量。當(dāng)進(jìn)行多個(gè)變量變異程度的比較時(shí),若各變量平均值相差較大,直接使用標(biāo)準(zhǔn)差進(jìn)行變異程度度量效果不好,則需要采用變異系數(shù)來(lái)比較。19個(gè)樣地植物物種重要值均值差異較大,因此重要值變異程度需通過(guò)變異系數(shù)來(lái)反映。計(jì)算公式[9]如下:
Vc=Ds/Nm×100%。
(2)
式(2)中,Vc為植物重要值變異系數(shù);Ds為10種主要植物在19個(gè)樣地中重要值的標(biāo)準(zhǔn)差;Nm為平均值。
(2)資源環(huán)境梯度的劃分
樣地土壤體積含水量(以下簡(jiǎn)稱(chēng)土壤含水量)、電導(dǎo)率以野外測(cè)定的每個(gè)樣地的5個(gè)樣方平均值為準(zhǔn);實(shí)驗(yàn)室測(cè)定19份土樣土壤有機(jī)質(zhì)含量、pH值,用GPS記錄樣地經(jīng)緯度,并用GIS軟件提取樣地高程數(shù)據(jù);共得到19個(gè)樣地的土壤含水量、電導(dǎo)率、有機(jī)質(zhì)含量、pH值以及高程數(shù)據(jù)。在上述5個(gè)環(huán)境因子上,分別以等距離的梯度間隔將其從小到大劃分為10個(gè)等級(jí)[10],作為資源環(huán)境梯度標(biāo)識(shí)(表1)。
表1 5種資源環(huán)境因子等級(jí)梯度劃分
Table 1 Position of the 5 resource-environmental factors on rank gradient
等級(jí)序號(hào)高程/mpH值電導(dǎo)率/(μS·cm-1)土壤含水量/%土壤有機(jī)質(zhì)含量/(g·kg-1)113.80~13.974.52~4.6117.40~23.024.88~5.931.35~1.86213.97~14.134.61~4.7123.02~28.645.93~6.971.86~2.38314.13~14.294.71~4.8028.64~34.266.97~8.022.38~2.89414.29~14.464.80~4.8934.26~39.888.02~9.072.89~3.40514.40~14.624.89~4.9939.88~45.509.07~10.123.40~3.92614.62~14.794.99~5.0845.50~51.1210.12~11.163.92~4.43714.79~14.955.08~5.1751.12~56.7411.16~12.214.43~4.94814.95~15.125.17~5.2656.74~62.3612.21~13.264.94~5.45915.12~15.285.26~5.3662.36~67.9813.26~14.305.45~5.971015.28~15.455.36~5.4567.98~73.6014.30~15.355.97~6.48
(3)生態(tài)位寬度測(cè)定
利用Levins生態(tài)位寬度公式計(jì)算研究區(qū)10種主要植物在5種資源環(huán)境因子梯度上的生態(tài)位寬度[11]:
(3)
式(3)中,Bi為植物物種i的生態(tài)位寬度;r為各資源環(huán)境因子等級(jí)數(shù);nij為物種i在第j個(gè)資源環(huán)境等級(jí)中的重要值(以物種i在資源環(huán)境等級(jí)j上的重要值的平均值為準(zhǔn));Ni為物種i在所有資源環(huán)境等級(jí)中重要值的總和。
(4)生態(tài)位重疊計(jì)算
利用Pianka生態(tài)位重疊公式計(jì)算研究區(qū)10種主要植物在5種資源環(huán)境因子梯度上的生態(tài)位重疊[12-13]:
(4)
式(4)中,Oik為植物物種i和k的生態(tài)位重疊值;r為各資源環(huán)境因子等級(jí)數(shù);nij和nkj為物種i和k在資源環(huán)境等級(jí)j上的重要值(以物種i和k在資源環(huán)境等級(jí)j上的重要值的平均值為準(zhǔn))。
2.1 植物物種重要值及其變異系數(shù)
2.2 研究區(qū)10種主要植物生態(tài)位寬度
表2 恒湖農(nóng)場(chǎng)茶葉港草洲10種主要植物的重要值和頻度
Table 2 Important values and frequencies of the 10 main plants in Chayegang Grassland of the Henghu Farm
種號(hào)物種重要值均值重要值標(biāo)準(zhǔn)差重要值變異系數(shù)/%頻度/%1灰化薹草(Carexcinerascens)0.4430.17840.394.742水田碎米薺(Cardaminelyra-ta)0.0460.069148.742.113蔞蒿(Artemisiaselengensis)0.0760.131170.842.114草(Phalarisarundinacea)0.2200.18182.378.955南荻(Triarrhenalutariori-paria)0.1410.221156.742.116藨草(Scirpustriqueter)0.0150.036232.515.797剛毛荸薺(Eleocharisvalleculosa)0.0060.019303.810.538旋鱗莎草(Cyperusmichelia-nus)0.0050.016318.410.539廣州蔊菜(Rorippacantonien-sis)0.0060.020331.610.5310蘆葦(Phragmitesaustralis)0.0340.110325.010.53
表3 5個(gè)因子梯度上10種主要植物生態(tài)位寬度
Table 3 Niche breadth of the 10 main plants along the 5-factor gradients
物種生態(tài)位寬度高程土壤pH值土壤電導(dǎo)率土壤含水量土壤有機(jī)質(zhì)含量總平均值灰化薹草(Carexcinerascens)8.546.618.168.969.128.28水田碎米薺(Cardaminelyrata)4.014.163.594.153.853.95蔞蒿(Artemisiaselengensis)3.392.843.134.493.973.56草(Phalarisarundinacea)7.615.825.555.947.246.43南荻(Triarrhenalutarioripar-ia)2.743.663.753.934.013.62藨草(Scirpustriqueter)1.002.351.912.702.782.15剛毛荸薺(Eleocharisvalleculo-sa)1.871.521.521.861.521.66旋鱗莎草(Cyperusmichelia-nus)1.981.711.711.981.981.87廣州蔊菜(Rorippacantonien-sis)2.001.581.581.911.911.80蘆葦(Phragmitesaustralis)1.511.641.951.641.641.68總平均值3.463.193.293.763.80
2.3 研究區(qū)10種主要植物45個(gè)種對(duì)生態(tài)位重疊
表4 10種主要植物45個(gè)物種對(duì)在高程和土壤pH值因子梯度上的生態(tài)位重疊
Table 4 Niche overlaps of 45 species pairs of the 10 main plants along the elevation and soil pH gradients
物種1234567891010.700.460.940.520.300.450.290.300.3220.590.160.680.030.030.630.210.200.1130.470.220.300.650.560.050.640.580.1240.630.810.270.300.070.530.370.390.4850.720.190.690.370.780.000.000.000.1060.710.180.090.310.390.000.000.000.0070.100.580.000.810.000.000.320.360.4980.070.450.250.730.000.000.890.990.6190.060.440.200.740.000.000.921.000.69100.510.000.180.290.320.390.000.000.00
表5 10種主要植物45個(gè)物種對(duì)在土壤電導(dǎo)率和土壤含水量因子梯度上的生態(tài)位重疊
Table 5 Niche overlaps of 45 species pairs of the 10 main plants along the soil conductivity and water content gradients
物種1234567891010.620.460.760.540.570.240.230.210.4920.670.050.680.110.100.550.450.460.4530.540.150.110.920.150.010.110.090.1840.690.680.240.130.290.590.620.620.2450.530.040.730.150.280.050.010.010.1660.490.150.320.150.620.000.000.000.0070.260.530.000.810.030.000.890.920.2280.200.300.360.640.030.000.671.000.2290.190.290.310.680.030.000.730.990.18100.570.310.050.300.010.000.000.000.00
表6 10種主要植物45個(gè)物種對(duì)在土壤有機(jī)質(zhì)含量因子梯度上的生態(tài)位重疊
Table 6 Niche overlaps of 45 species pairs of the 10 main plants along the soil organic matter gradient
物種12345678920.6830.460.3540.830.350.3550.530.260.640.3260.410.430.360.490.5270.280.330.030.490.120.7980.260.340.440.420.070.560.7490.260.330.380.440.080.610.810.99100.340.000.120.200.540.000.000.000.00
2.4 5個(gè)因子梯度上植物種群生態(tài)位寬度與重要值及其變異系數(shù)的相關(guān)程度
通過(guò)Spearman秩相關(guān)分析檢驗(yàn)不同因子梯度上植物種群生態(tài)位寬度分別與重要值、重要值變異系數(shù)之間的相關(guān)程度(圖1)。
*表示相關(guān)顯著(P<0.05),**表示相關(guān)極顯著(P<0.05)。
由圖1可知,土壤pH值、土壤電導(dǎo)率、含水量、有機(jī)質(zhì)含量因子梯度上10種植物生態(tài)位寬度與重要值之間均呈極顯著正相關(guān)(P<0.01),高程因子梯度上兩者呈顯著正相關(guān)(P<0.05),相關(guān)系數(shù)r分別為0.855、0.939、0.794、0.855和0.733。而在上述5個(gè)因子梯度上10種植物生態(tài)位寬度與重要值變異系數(shù)之間均呈極顯著負(fù)相關(guān)(P<0.01),r分別為-0.927、-0.867、-0.903、-0.879和-0.782。
物種重要值和種群生態(tài)位是衡量植物在群落中地位與作用的2個(gè)指標(biāo),但所代表的意義完全不同。重要值反映的是某個(gè)植物種群在群落中的相對(duì)重要性,而生態(tài)位寬度揭示的是物種對(duì)環(huán)境的適應(yīng)性及資源利用能力。筆者研究中,5個(gè)因子梯度上植物種群生態(tài)位寬度與重要值之間均呈正相關(guān),且達(dá)顯著(P<0.05)或極顯著水平(P<0.01),這與潘高等[25]探究南方紅壤丘陵區(qū)3種森林群落內(nèi)主要草本植物種群生態(tài)位特征得出的結(jié)果較一致。不過(guò)兩者之間未必存在必然聯(lián)系,筆者研究中蘆葦?shù)闹匾荡笥谒懖?但在土壤pH值、含水量和有機(jī)質(zhì)含量梯度上,前者的生態(tài)位寬度卻小于后者,這是因?yàn)橹匾挡⒎鞘怯绊懛N群生態(tài)位寬度的唯一因素。錢(qián)逸凡等[26]研究認(rèn)為分布頻度也是決定物種生態(tài)位寬度的主要因素,物種分布頻度越高,其生態(tài)位寬度越寬。筆者研究中,物種重要值變異系數(shù)也是影響物種生態(tài)位寬度的重要因素,它們之間呈負(fù)相關(guān),此與前人研究結(jié)論[27]一致。研究區(qū)植物群落中雖然蔞蒿重要值大于水田碎米薺,但其重要值變異系數(shù)較后者大,因此在高程、土壤pH值、電導(dǎo)率3個(gè)環(huán)境梯度上生態(tài)位寬度均小于后者。
種群生態(tài)位研究對(duì)于了解植物在群落中的功能地位、生態(tài)適應(yīng)性以及生態(tài)相似性方面具有重要作用,為進(jìn)一步研究植被分布特征和生物多樣性形成機(jī)制提供了科學(xué)依據(jù)[23]。需要注意的是,筆者基于5個(gè)資源環(huán)境梯度,對(duì)植物種群生態(tài)位寬度及生態(tài)位重疊的計(jì)算具有一定的時(shí)間和空間限制[28]。首先,隨著時(shí)間的推移,植物群落中的環(huán)境因子以及優(yōu)勢(shì)種會(huì)發(fā)生改變,生態(tài)位寬度與環(huán)境因子之間關(guān)系密切,當(dāng)環(huán)境條件發(fā)生變化時(shí),植物對(duì)生態(tài)環(huán)境適應(yīng)性以及資源利用能力也會(huì)隨之改變;而當(dāng)群落優(yōu)勢(shì)種發(fā)生變化時(shí),不同物種之間生態(tài)位重疊值也會(huì)有所不同。其次,當(dāng)研究尺度不同時(shí),群落樣地環(huán)境以及不同樣地環(huán)境之間的相關(guān)性或者空間自相關(guān)也會(huì)出現(xiàn)差異,間接導(dǎo)致植物種群生態(tài)位寬度、物種間生態(tài)位重疊值發(fā)生變化。
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(責(zé)任編輯: 李祥敏)
Niches of the Major Plant Populations in Grasslands Typical of the Poyang Lake Wetland in Five Resources-Environmental Gradients.
DUANHou-lang1,2,ZHAOAn1,2,YAOZhong3
(1.Key Laboratory of Poyang Lake Wetland and Watershed Research, Ministry of Education, Jiangxi Normal University, Nanchang 330022, China;2.School of Geography and Environmental Sciences, Jiangxi Normal University, Nanchang 330022, China;3.Jiangxi Academy of Sciences, Nanchang 330096, China)
Data of 10 dominant plant species and 5 resources-environmental factors (elevation, soil pH, electrical conductivity, water content and organic matter content) were gathered through surveys of 95 quadrates (1 m×1 m) in 19 sample plots (5 m×5 m) in the Chayegang Grassland of the Henghu Farm, typical of the Poyang Lake wetland, and analyzed for breadths and overlaps of the ecological niches of the plant populations, using the indices of Levins niche breadth and Pianka niche overlap. Results show that: (1)CarexcinerascensandPhalarisarundinaceaare the dominant species in the region and have broader niche along the five resources-environmental gradients, whileCyperusmichelianus,Rorippacantoniensis,Phragmitesaustralisand some others are accidental species and have relatively narrow niches; (2) The 10 plant species rarely overlap in niche, but exception exists withEleocharisvalleculosa,CyperusmichelianusandRorippacantoniensis, which overlap each other much more than the other plant species pairs; (3) Of the 10 major plant populations, niche breadth is significantly (P<0.05) or extra-significantly (P<0.01) and positively related to other important values, but extra-significantly (P<0.01) and negatively related to variation coefficient of their important values. The 10 dominant plant species vary sharply in resource-environment utilization capability and adaptability to the 5 environmental factors, and are generally low in niche overlap. Importance values of the plant species are the major factors dictating niche breadth.
Poyang Lake wetland; plant population; niche breadth; niche overlap
2016-07-04
國(guó)家自然科學(xué)基金(81260449);江西省青年科學(xué)基金(201232BAB214022);江西省教育廳2012年度科技項(xiàng)目(CJJ12185)
Q948
A
1673-4831(2017)03-0225-09
10.11934/j.issn.1673-4831.2017.03.005
段后浪(1992—),男,安徽六安人,碩士生,主要從事植被生態(tài)學(xué)方面的研究。E-mail: duanhl2408@126.com
① 通信作者E-mail: zhaoanjxsd@126.com