第 42 卷 第 6 期 水 生 生 物 学 报 Vol. 42, No. 6

2018 年 11 月 ACTA HYDROBIOLOGICA SINICA Nov., 2018 doi: 10.7541/2018.141

MAPPING SPATIOTEMPORAL TRENDS IN THE ABUNDANCE AND DISTRIBUTION OF MACROPHYTES IN HONGZE

GUO Chuan-Bo1, LI Wei1, ZHANG Ying-Xue1, 2, XIA Wen-Tong1, 2, XIN Wei1, CHEN Yu-Shun1, 2 and LI Zhong-Jie1, 2 (1. State Key Laboratory of Freshwater Ecology and Biotechnology, Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan 430072, ; 2. University of Chinese Academy of Sciences, Beijing 100049, China)

Abstract: A comprehensive investigation on macrophyte community in was conducted seaso- nally from May 2010 to February 2011. Overall, twelve species representing eight families of macrophytes were identified in Hongze Lake, including nine species of submerged plants, two species of floating-leaved plants, and one species of emerging plant. In general, Potamogeton malaianus, P. maackianu, P. pectinatus and P. crispus were the four dominant species throughout the whole year, the highest biomass of macro- phytes was presented in autumn, followed by summer and winter, while spring had the lowest biomass of macrophytes. Based on field data, we used kriging interpolation in ArcGis to map the spatiotemporal distribu- tion of the entire macrophyte community as well as each of the four dominant species. From the GIS maps we observed that the northern area of the lake, namely the Chengzihu region, had the highest biomass of macro- phytes potentially as a result of better water quality and greater transparency. Potential factors that affected the community structure, biomass, and distribution patterns of macrophytes considerably were then discussed. The results of this study illuminate the need for more information on the role and importance of aquatic mac- rophytes in shallow lake ecosystems. Conservation of macrophytes should be taken to maintain the lake eco- system health.

Key words: Macrophyte community; GIS and GPS; Hongze Lake; Spatiotemporal patterns; Shallow CLC number: S932.8 Document code: A Article ID: 1000-3207(2018)06-1153-10

Macrophytes play an important role in the struc- macrophytes in shallow lake ecosystems is related to ture and functioning of shallow freshwater ecosys- their structural attributes such as species composition, tems[1—3]. They serve as a base of most aquatic food- distribution, abundance, and diversity. These attri- webs, and also produce food for biota, provide refu- butes which in turn rely on various environmental gia for other organisms[4], anchor soft bottom sedi- factors including water level, water temperature, sub- ments, remove suspended particles and nutrients[4], strate composition, disturbance, competitive interac- and contribute to the promotion and maintenance of tions, herbivory, epiphyte loading, water quality and aquatic foodwebs and ecosystem services[5, 6]. Macro- sediment nutrients[3, 9—11]. Therefore understanding phytes also contribute to the general fitness and di- and quantifying the community structure and spati- versity of aquatic ecosystems by serving as indicators otemporal patterns of macrophytes is indispensable for water quality and aiding in nutrient cycling[7, 8]. As for integrated management practices and aquatic con- a result of their significance, macrophytes are com- servation in shallow lake ecosystems. An example of monly viewed as one of the most important foci in a macrophyte-driven shallow water ecosystem is that shallow lake ecosystems. Generally, the function of of Hongze Lake.

Received date: 2016-06-21; Accepted date: 2018-01-17 Foundation item: Supported by Chinese Academy of Sciences Grants (Y62302, Y45Z04,Y55Z06 and QYZDB-SSW-SMC041); National Natural Science Foundation of China (31602158) Corresponding author: Chen Yu-Shun, E-mail: [email protected] 1154 水 生 生 物 学 报 42 卷

Hongze Lake (33°06′—33°40′N, 118°10′— 118° ecosystem provides multiple benefits to various stake- 52′E), which lies in the middle reach of the Huaihe holders. Hongze Lake acts as a navigation junction, River, is the fourth largest freshwater lake in China aids flood-and-drought resistance, provides fishing 2 with a surface area of 1597 km , a mean water depth and aquaculture opportunities, as well as supports [12] of 1.7 m . Hongze Lake is also the largest storing biodiversity conservation and tourism in eastern reservoir and water channel on the east route of the China[12, 13]. China’s South-North Water Division Project (SN- 1.2 Sampling procedure [13] WDP) . Hongze Lake ecosystems provide multiple Four field surveys were conducted in Hongze benefits that are fundamental to human wellbeing, in- Lake during 2010－2011 in spring (May), summer cluding commercial navigation, flood control, com- (August), autumn (November) and winter (February) mercial fishing, aquaculture, biodiversity conserva- respectively. Finally, a total of 631 sites, each consist- [12, 13] tion, and tourism in eastern China . However, ing of a 0.125 m2 quadrat, were randomly selected in Hongze Lake has also experienced numerous natural the whole lake according to the lake morphology and and artificial disturbances in recent years, such as re- habitat heterogeneity. Seasonaly, we sampled 173, gional climate change, high nutrients inputs, water 173, 130 and 155 sites during spring, summer, au- conservancy construction, recreational and commer- tumn and winter respectively. Throughout sampling, cial fishing, and aquaculture. As a result of these dis- we avoided selection of area near intensive block nets turbances, aquatic macrophytes have seen great de- in the lake (Fig. 1). Within each 0.125 m2 sampling clines, specifically the biomass and species diversity [13, 14] quadra in each of the sampling site, all macrophytes has decreased whilst distributional areas shrunk . were uprooted with a bottom sampler. All macro- These alterations have the potential to affect the dis- phytes were then cleaned, identified into species fol- tribution patterns and community structure of other lowing Cook[18] and then weighed to the nearest 0.1 g organisms which can ultimately, affect the overall [15] wet weight in the field immediately. The geographic health of the lake ecosystem . coordinates of each sampling site were recorded us- However, very little knowledge about the spatial ing a handling GPS unit (Garmin eTrex 301). and temporal dynamics of macrophytes in Hongze 1.3 Data processing and GIS mapping Lake exists, mainly due to the fact that large-scale and The relative biomass (B ), frequency of occur- long-term field monitoring efforts for macrophytes in ri rence (F ) and degree of dominance (D ) were calcu- such a large lake presents numerous challenges. Pre- i i lated according to the following equations[19]: vious studies of macrophytes in Hongze Lake have only been conducted with very limited datasets either B ri = B i=B t 100 (1) [14, 16] £ on a small spatial scale or during a single season . Fi = N i=N t 100 (2) The current study was conducted by integrating GPS £ D i = (B ri + Fi) =2 (3) and GIS technologies with field monitoring data to simulate the community structure and spatiotemporal Where Bri is the relative biomass of the species i, Bi is patterns and dynamics of macrophytes in the whole the biomass of the species i, Bt is the total biomass of Hongze Lake. The main objective of our study was to all the species; Fi is the frequency of occurrence of map the distribution patterns and seasonal dynamics species i, Ni and Nt is the occurrence number of spe- of macrophytes community structure in Hongze Lake cies i and the total sampling sites, respectively; Di is across broad spatial and temporal scales. The results the degree of dominance. of this study will benefit the conservation of macro- The spatiotemporal patterns of the total macro- phytes resources in not only Hongze but other shal- phytes biomass and four dominant macrophyte were low lakes as well. then mapped using Arcgis 10.1 (ESRI) using the Kri- ging interpolation utility on a map of Hongze Lake 1 Materials and Methods that was modified from Google Earth (Google 1.1 Study area Earthtm). Hongze Lake is the fourth largest freshwater lake 2 2 Results in China with a surface area of 1597 km , and a mean water depth of 1.7 m[12]. The rivers draining into 2.1 Community structure of macrophytes Hongze Lake are located primarily along the western A total of twelve species belong to eight fami- bank of the lake. Among them, the Huaihe River is lies of macrophytes were identified from the whole the largest, and contributes 87% of the total inflow to Hongze Lake. Collections included nine species of the lake[12]. Hongze Lake is a transitional lake, mean- submerged plants (Potamogeton malaianus, Myrio- ing the water levels of the lake often undergo large phyllum spicatum, P. maackianus, P. pectinatus, Hy- annual and seasonal changes[17]. The Hongze Lake drilla verticillata, Ceratophyllum demersum, Elodea 6 期 郭传波等: 洪泽湖大型水生植物群落结构和时空格局的GIS模拟 1155 nuttallii, P. crispus, Vallisneria natans), two species lowest macrophyte biomass (Fig. 2). In particular, of floating-leaved plants (Eichhornia crassipes, high biomasses of P. crispus observed mostly during Trapa bispinosa) and one species of emerging plant spring and winter had disappeared by summer, while (Miscanthus sacchariflorus). The greatest biomass of biomasses of E. nuttallii and M. sacchariflorus in- macrophytes were observed during autumn, followed creased only during spring (Tab. 1; Fig. 2). by summer and winter, while spring contained the Macrophyte communities in Hongze Lake were

118°20′E 118°30′E 118°40′E 118°50′E

33°40′N SpringN Summer N

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0 5 10 20 Sampling site 0 5 10 20 Sampling site km km Autumn N Winter N

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Sampling site Sampling site 0 5 10 20 0 5 10 20 km km Fig. 1 Location and distribution of sampling sites in four seasons in Hongze Lake

40 V. Natans 35 T. Bispinosa M. Sacchariflorus

) 30 2 P. Crispus

g/m 25

4 E. Nuttallii E. Crassipes 20 C. demersum 15 H. verticillata

Biomass (×10 10 P. Pectinatus P. Maackianus 5 M. Spicatum 0 P. Malaianus Spring Summer Autumn Winter Fig. 2 Seasonal variations of species composition of macrophytes community in Hongze Lake 1156 水 生 生 物 学 报 42 卷 dominated by P. malaianus, P. maackianu, P. pec- and northeastern part of the lake in autumn, though tinatus and P. crispus throughout the year (Fig. 2). biomasses were never as high as those observed dur- The dominant species showed significant difference ing summer or winter (Fig. 5). P. maackianus bio- among the four seasons: P. maackianus and P. cris- mass was elevated in the northern part of the lake dur- pus dominated the macrophyte community during ing spring and autumn, but was limited to the western

Spring with Di values of 32.1% and 20.1%, respec- and central parts of the lake during winter (Fig. 6). tively. During Summer, communities shifted more to- Generally, significant high biomass patch can be also wards P. malaianus (21.3%), C. demersum (15.1%), seen in the central and west portion of the lake in P. pectinatus (9.3%) and T. bispinosa (8.9%) which spring and winter. During summer, P. crispus could dominated the community. By autumn, communities not be detected in the lake (Fig. 7). had transitioned over to P. malaianus (28. 5%) and P. 3 Discussions maackianus (23.7%). And P. maackianus (18.7%), P. 3.1 Community structure of macrophytes malaianus (18.3%) and P. pectinatus (13.7%) domina- The biomass of macrophytes was historically ting the macrophyte communities in winter (Tab. 1). high in Hongze Lake prior to the construction of a 2.2 Spatiotemporal distribution patterns of macro- dam between Hongze Lake and Huaihe River[16]. Sig- phytes nificant degradation of macrophytes communities was Using the GIS simulation, the spatiotemporal observed following dam construction, such that both distributions of macrophyte biomass was mapped in a biomass and diversity were decreased. In 1960, there set of quadruple lattices. The greatest biomass of were 28 species of macrophytes were recorded, com- macrophytes was distributed in the northern part of pared to 29 species in 1981[20]. However, Liu, et [16] the Hongze Lake, where large areas of macrophytes al. reported 25 species of macrophyte species in the existed during summer, and smaller patch of high bio- early 2000s. All of these figures sharply contrast with mass was found in autumn (Fig. 3). the current study, where only 12 species were recor- Distribution patterns of P. malaianus varied sig- ded. Although these suggest that macrophyte di- nificantly across seasons (Fig. 4). P. malaianus had versity in Hongze Lake has declined sharply, results lowest biomass in spring and highest in autumn (Fig. 4). also partly variations in species collection area and sampling methods used across studies. In the current Although the western, northern, and northeastern study, we focused on macrophytes in the open-water parts of Hongze Lake were occupied by P. malaianus areas that excluded littoral zones and border wetlands in summer and autumn, distributions were limited to that would likely support more species. In addition, the central and northeastern parts of the lake by there was a large area in the lake being used for en- winter (Fig. 4). High-biomass patches of P. pec- closure aquaculture, which was not accessible to our tinatus were found in the northern and northeastern sampling. Thus, our current estimate of 12 macro- part of Hongze Lake during summer, but were limi- phyte species is likely an underestimate. Liu, et al.[16] ted to only the northern part of the lake in winter (Fig. 5). revealed that P. malaianus was the dominant species P. pectinatus biomasses also were high in the western in Hongze Lake during their sampling in the early

Tab. 1 Seasonal variation of macrophyte quantitative characteristics in Hongze Lake Spring Summer Autumn Winter Species Bri (%) Fi (%) Di (%) Bi (%) Fi (%) Di (%) Bi (%) Fi (%) Di (%) Bi (%) Fi (%) Di (%) P. malaianus 0.3 1.1 0.7 22.9 10.6 21.3 37.2 19.7 28.5 27.1 9.6 18.3 M. spicatum 3.6 2.3 2.9 1.3 3.03 5.6 3.8 9.8 6.8 0 0 0.00 P. maackianus 58.4 5.7 32.1 6.7 0.00 6.2 41.5 5.8 23.7 32.3 5.1 18.7 P. pectinatus 1.1 1.1 1.1 14.6 4.6 9.3 9.0 4.1 6.5 22.8 4.5 13.7 H. verticillata 0.00 0.00 0.00 0.08 2.3 2.6 5.0 5.2 5.1 0 0 0.00 C. demersum 0.01 0.6 0.3 24.5 6.8 15.1 0.8 5.8 3.3 0.8 0.6 0.7 E. crassipes 2.3 2.3 2.3 11.7 2.3 7.0 1.8 2.3 2.1 2.1 0.6 1.4 E. nuttallii 1.9 3.4 2.7 0 0 0 0 0 0 0 0 0 P. crispus 31.7 8.6 20.1 0 0 1.2 0.1 2.3 1.2 12.3 1.9 7.1 M. sacchariflorus 0.6 0.6 0.6 0 0 0 0 0 0 0 0 0 T. bispinosa 0.04 0.6 0.3 17.9 6.1 8.9 0 0 0 0 0 0 V. natans 0.00 0.00 0.00 0.4 1.5 1.4 0.6 2.3 1.5 2.7 1.9 2.3 6 期 郭传波等: 洪泽湖大型水生植物群落结构和时空格局的GIS模拟 1157

118°20′E 118°30′E 118°40′E 118°50′E 33°40′N SpringNN Summer

Hongze Lake Hongze Lake 33°30′N

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0 0 0 5 10 20km 0 5 10 20km Fig. 3 Distribution of total biomass (g/m2) of macrophytes in Hongze Lake by GIS simulation using kriging interpolation utility

118°20′E 118°30′E 118°40′E 118°50′E 33°40′N SpringNN Summer

Hongze Lake Hongze Lake 33°30′N

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Hongze Lake Hongze Lake

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0 0 0 5 10 20km 0 5 10 20km Fig. 4 Distribution of the biomass (g/m2) of P. malaianus in Hongze Lake by GIS simulation using kriging interpolation utility 1158 水 生 生 物 学 报 42 卷

118°20′E 118°30′E 118°40′E 118°50′E 33°40′N SpringNN Summer

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0 0 0 5 10 20km 0 5 10 20km Fig. 5 Distribution of the biomass (g/m2) of P. pectinatus in Hongze Lake by GIS simulation using kriging interpolation utility

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Hongze Lake Hongze Lake 33°30′N

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Hongze Lake Hongze Lake

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0 0 0 5 10 20km 0 5 10 20km Fig. 6 Distribution of the biomass (g/m2) of P. maackianus in Hongze Lake by GIS simulation using kriging interpolation utility 6 期 郭传波等: 洪泽湖大型水生植物群落结构和时空格局的GIS模拟 1159

118°20′E 118°30′E 118°40′E 118°50′E

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Hongze Lake Hongze Lake 33°30′N

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Hongze Lake Hongze Lake

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0 5 10 20 0 0 5 10 20 0 km km

Fig. 7 Distribution of the biomass (g/m2) of P. crispus in Hongze Lake by GIS simulation using kriging interpolation utility

2000s, which was consistent with our current find- 3.2 Possible factors affecting distribution patterns ings. However, we also argue that the dominant spe- There is ample evidence in the literature that cies exhibited significant seasonal variation. For exam- many factors have affected the distribution, and spe- ple, P. maackianus and P. crispus dominated the mac- cies composition of macrophyte communities. These rophyte community in spring, P. malaianus, C. de- factors include light, water temperature, water quality mersum, P. pectinatus and T. bispinosa dominated the changes and nutrient enrichment, sediment composi- summer community, P. malaianus and P. maackia- tion, and water level fluctuations[22—24]. Light and nus dominated the autumn community, P. maackia- temperature within the context of water depth, season, nus, P. malaianus and P. Pectinatus dominated the and latitude are of paramount importance in dictating winter community. Overall, P. malaianus, P. maackia- macrophyte distributions, thereby indirectly influen- nu, P. pectinatus and P. crispus were the four most cing lake productivity and macrophyte species com- dominant species in Hongze Lake. Some of the sea- position[25]. The distribution patterns of macrophytes sonal variation was due to basic life histories. For in- are largely driven by water clarity, which affects the stance, P. crispus was only observed in spring and amount of light reaching the lake bottom where plants winter because this species dies off gradually during root[26—28]. Water quality changes and associated nu- spring and summer, but will sprout and grow during trient enrichment often cause considerable variation in winter[21]. the species richness, composition, and density for a 1160 水 生 生 物 学 报 42 卷 variety of aquatic vegetation species[3, 29, 30]. In the cur- bution and ecological status of the Turiec River (Slovakia): rent study, the northern and northeastern parts of changes after seven years [J]. Archives of Biology Sciences, Hongze Lake functioned like a “closed” system 2009, 61(2): 297—306 (Chengzihu area) in that water quality was in gene- [3] Tamire G, Mengistou S. Macrophyte species composition, [31] rally good condition with high transparency . Not distribution and diversity in relation to some physicochemi- surprisingly, more macrophyte species that exhibited cal factors in the littoral zone of Lake Ziway, Ethiopia [J]. high biomasses throughout the whole year were dis- African Journal of Ecology, 2012, 51(1): 66—77 tributed in this area of the lake. However, in the cent- ral and southern parts of Hongze Lake, high nutrient [4] Madsen J D, Bloomfield J A, Sutherland J W, et al. 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The effect of temperature and day fect lake limnology, which in turn, affects species length on the germination of Potamogetonnodosus tubers [J]. zonation, distribution, biomasses, and richness of Journal of Freshwater Ecology, 1995, 10: 125—128 [34] macrophytes on decadal time scales . Hongze Lake [8] Carpenter S R, Lodge D M. Effects of submersed macro- formed as a reservoir in the 1950s when Sanhe Dam phytes on ecosystem processes [J]. Aquatic Botany, 1986, was constructed between the lake and the Huaihe 26(3-4): 341—370 River. As a result, base water levels have increased [9] Jaikumar M, Chellaiyan D, Kanagu L, Kumar P S, Stella C. [16] from 10.6 to 12.4 m during the time ; by the 1990s, Distribution and succession of aquatic macrophytes in Chilka water levels were further increased to 13.0 m[35]. The Lake –India [J]. 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洪泽湖大型水生植物群落结构和时空格局的GIS模拟

郭传波1 李 为1 张映雪1, 2 夏文彤1, 2 辛 未1 陈宇顺1, 2 李钟杰1, 2 (1. 中国科学院水生生物研究所，淡水生态与生物技术国家重点实验室, 武汉 430072; 2. 中国科学院大学, 北京 100049)

摘要: 2010—2011年对洪泽湖大型水生植物进行了4个季度全面的调查和研究, 共发现大型水生植物8科12种, 其中沉水植物9种, 挺水植物1种, 浮叶植物2种。马来眼子菜(Potamogeton malaianus)、微齿眼子菜(P. maackia- nu)、篦齿眼子菜(P. pectinatus)和菹草(P. crispus)为全年优势度较高的水生植物, 但4个季节大型水生植物的 优势种类组成差异明显。秋季的水草生物量最高, 其次为夏季和冬季, 春季最低。结合GPS (Global Position System)和GIS (Geographic Information System), 利用GIS的Kring插值法对洪泽湖大型水生植物总生物量及主 要优势物种的时空分布进行了可视化模拟。结果发现洪泽湖现阶段大型水生植物分布区域主要集中在湖区 北部水质较好、透明度较高且相对封闭的成子湖区。文章也分析了洪泽湖大型水生植物变迁的潜在影响因 子, 为水生植物保护和生态系统健康提供了基础依据。

关键词: 大型水生植物; GIS和GPS; 洪泽湖; 时空格局; 浅水湖泊

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主 编 Chief Editor 桂建芳 GUI Jian-Fang 副主编 Associate Editor 解绶启 XIE Shou-Qi 委 员 Members (以姓氏拼音为序) 蔡庆华 CAI Qing-Hua 曹文宣 CAO Wen-Xuan 常剑波 CHANG Jian-Bo 陈家宽 CHEN Jia-Kuan 陈宜瑜 CHEN Yi-Yu 陈毅锋 CHEN Yi-Feng 高坤山 GAO Kun-Shan 何舜平 HE Shun-Ping 胡征宇 HU Zheng-Yu 李文鑫 LI Wen-Xin 李钟杰 LI Zhong-Jie 林浩然 LIN Hao-Ran 刘永定 LIU Yong-Ding 麦康森 Mai Kang-Sen 聂 品 NIE Pin 曲久辉 QU Jiu-Hui 宋立荣 SONG Li-Rong 唐启升 TANG Qi-Sheng 王 丁 WANG Ding 吴灶和 WU Zao-He 吴振斌 WU Zhen-Bin 相建海 XIANG Jian-Hai 肖 伟 XIAO Wei 谢 平 XIE Ping 谢小军 XIE Xiao-Jun 熊邦喜 XIONG Bang-Xi 熊思岳 XIONG Si-Yue 徐旭东 XU Xu-Dong 杨先乐 YANG Xian-Le 于 丹 YU Dan 余其兴 YU Qi-Xing 游 力 YOU Li 张奇亚 ZHANG Qi-Ya 朱作言 ZHU Zuo-Yan Harald Rosenthal (德国) 编辑部 Editorial office 杜新征 Du Xin-Zheng 余 茜 YU Xi 叶文娟 YE Wen-Juan