Threshold value of Benthic Pollution Index (BPI) for a muddy healthy benthic faunal community and its application to Jinhae Bay in the southern coast of Korea

For the determination of benthic community health criteria of a biotic index, Benthic Pollution Index (BPI), the faunal data on macrobenthic community of Deukryang Bay collected in 2012 were used. Each macrobenthic fauna was classified into 4 functional groups according to their feeding and life history strategies and BPI was calculated using the abundance of fauna for these functional groups. Amphipods were the dominant faunal group belonging to Functional Group (FG) II in Deukryang Bay during all seasons. The BPI value fluctuated seasonally from 55 to 61, and the overall mean value of BPI in the reference area was 60, which was estimated as the threshold value for healthy communities. We tried to use BPI as an assessment tool for macrofaunal community health status by categorizing the BPI values into 5 grades. If the BPI value of the community is more than 60, the health status of a macrobenthic community will be assessed as ‘Excellent’ (Grade 1) and as ‘Good’ (Grade 2), if between 40 and 60, as ‘Fair’ (Grade 3), if between 30 and 40, as ‘Poor’ (Grade 4), if between 20 and 30, and finally the health status of a community will be assessed as ‘Very Poor’ (Grade 5), if the BPI value is less than 20. This assessment tool using BPI was applied to macrobenthic communities in Jinhae Bay including the Special Management Area of South Korea. In Jinhae Bay, the FG IV containing a spionid species, Paraprionospio patiens was the most dominant group. The values of BPI in Jinhae Bay were very low and ranged from 11 to 23. The ecological health status of macrobenthic communities in Jinhae Bay was ‘Very Poor’ at more than 50% of sites, especially in summer.     

山东宁津附近海域大型底栖动物群落结构及季节变化

根据2009年11月至2010年8月在山东宁津附近海域进行4个季度月航次调查所得大型底栖动物采泥资料,运用优势度指数、物种多样性指数及Cluster聚类和MDS排序等方法,分析了该海域的大型底栖动物群落特征和季节变化。结果表明:4个航次在底泥样品中共鉴定大型底栖生物11门159种,其中,多毛类动物73种,是主要的优势类群,甲壳动物39种,软体动物25种,棘皮动物9种,其他类群共13种。主要优势种有拟特须虫、丝异蚓虫、索沙蚕、轮双眼钩虾等。宁津邻近海域年平均生物量和丰度分别为9.144g/m~2和233.90ind./m~2,均较低且存在季节性差异。丰富度指数(D)、均匀度指数(J)和多样性指数(H′)的年平均值分别为2.436、0.800和2.924,次级生产力较低,平均值仅为1.903g(AFDW)/(m~2·a)。聚类分析结果表明,秋、春航次群落分布格局均包含近岸群落和远岸群落,但各群落间界线不明显,群落结构受扰动情况不明显。     

Seasonal variations in the community structures of macrobenthic fauna and their health status in an estuarine bay,Gwangyang Bay in Korea

Macrobenthic fauna in an estuarine Gwangyang Bay, southern Korean coast, were investigated to uncover recent variations in their community structures. In the study area, macrobenthic faunal communities were mainly composed of polychaete worms which were the most abundant faunal group with the highest values in species number and density, while mollusks accounted for the highest proportion in total biomass. There was no clear seasonal difference in species richness during the two year period of the investigation, but the mean density and biomass increased every spring and summer due to the mass recruitment of Theora fragilis. The Shannon’s diversity index (H') was more than 2.0 during most sampling seasons and did not show any significant seasonal difference except for the data in August, 2011 when azoic conditions occurred. The community structures of macrobenthos in Gwangyang Bay did not show any remarkable change in the dominance of the two top dominant species, Scoletoma longifolia and Heteromastus filiformis, which abundantly occurred in all seasons, except for the abundance peaks associated with high occurrence of T. fragilis and Paraprionospio cordifolia, especially in spring and summer and in autumn, respectively. These fauna changes reflected the changes in the macrobenthic community health status in Gwangyang Bay, where stable conditions and a healthy status prevailed in winter, but a slightly disturbed status prevailed from spring to autumn.     

华南沿海海草床分布区大型底栖动物群落特征初探

为探究华南沿海海草床大型底栖动物群落特征、地区差异及其与海草群落间的关系,2020年在华南沿海12个海草床区域开展了大型底栖动物调查。共布设33个调查断面99个站位,在主要群落参数的基础上,运用相关性分析和聚类分析、nMDS排序、相似性百分比分析等群落生态学统计方法分析了底栖动物群落特征及其与海草群落的关系。共鉴定出大型底栖动物9大门类199种,其中,软体动物种类最多,占总种类数的39.7%。各区域底栖动物的平均栖息密度为155.7 ind/m²,青葛—龙湾(琼海)的平均栖息密度最低,珍珠湾(防城港)的平均栖息密度最高;平均生物量为118.36 g/m²,最低值出现在义丰溪(汕头),最高值出现在珍珠湾。各区域底栖动物的优势种以潮间带泥沙滩常见腹足类、双壳类和多毛类为主;底栖动物的种类多样性指数(H′)平均值为1.12,最低值出现在唐家湾(珠海),最高值出现在铁山港(北海);各调查区域底栖动物主要群落参数大致呈现“北部湾>海南沿岸>珠江口及粤东”的规律。华南沿海海草床分布区大型底栖动物群落的区域性特征较为明显,地理位置相近的海草床底...     

粤西海陵湾养殖区邻近海域大型底栖动物生态学特征

根据2014—2016年粤西海陵湾养殖区湾内和湾外邻近海域21°27′—21°38′N、111°42′—111°57′E 4个航次调查资料,对其大型底栖动物生态学特征进行了研究。结果表明,共鉴定大型底栖动物64种,春季种类最多为39种,秋季最低为17种。4季均以环节动物种类数最多,软体动物次之。不同季节间优势种有所差异,仅倍棘蛇尾Amphioplus sp.为周年优势种。春、夏、秋和冬季第一优势种分别为倍棘蛇尾、菲律宾蛤仔Ruditapes philippinarum、短吻铲荚螠Listriolobus brevirostris和平蛤蜊Mactra mera。平均丰度和生物量分别为213ind./m~2和15.4g/m~2,其中丰度以春季最高为248ind./m~2,夏季最低为167ind./m~2,而生物量则以秋季最高为28.0g/m~2,春季最低为2.4g/m~2;平面分布总体呈现湾外高于湾内的趋势。底栖动物Pielou均匀度和Shannon-Wiener多样性指数分别为0.95—1.00和1.56—4.07,其中时间分布规律明显,春季最高,秋季最低;空间上,两者无明显分布差异。群落结构时空差异显著,为沉积物类型、水深、底层无机氮和悬浮物以及捕食压力等因素共同影响的结果。     

桑沟湾大型底栖动物生物多样性研究

为了解桑沟湾大型底栖动物多样性现状,作者于2009年7月和12月对桑沟湾9个站位进行2个航次的采样,以种类组成、生物量和栖息密度为基础,采用物种多样性指数(Shannon-Wiener index)、物种丰富度指数(Species richnessindex)以及物种均匀度指数(Species evenness index)对大型底栖动物多样性及其影响因子进行了分析。结果表明:共采到大型底栖动物83种,其中多毛类45种,占种类总数的54.22%,软体动物11种(占13.25%),甲壳类23种(占27.71%),其他4种(占4.82%)。各站获得平均种类数在2～28种之间,种类最少的出现在夏季的C8站,最多的出现在夏季的C4站。不同养殖区生物多样性指数差异也很大,物种多样性(H′)和物种丰富度指数(D)均以贝类养殖区最高,藻类养殖区最低,而物种均匀度指数(J′)以贝藻混养区最高,藻类养殖区最低。生物多样性除了受种的数量及其个体密度的影响外,还与多种环境因素有关,相关性分析结果表明:水温、盐度、沉积物类型和硫化物含量是影响桑沟湾大型底栖动物生物多样性的主要环境因子。     

山东荣成临洛北湾夏季大型底栖动物群落生态学初步研究

利用2010年7月在山东荣成临洛北湾所获得大型底栖动物定量分析资料,研究了夏季该海域海带养殖区和邻近水域大型底栖动物在物种组成、生物量、丰度、优势种以及物种多样性等方面的特征和空间差异,并采用MDS和CLUSTER分析了大型底栖动物的群落结构,以期为该地区以后的科学研究提供基础数据.结果表明,本次调查通过采泥和拖网共采集到大型底栖动物88种,其中采泥得到大型底栖动物51种,拖网采得大型底栖动物45种,各站位的物种多样性差异明显,物种数量从2种到21种.与环境因子的相关性分析显示,各站的物种数和丰度与DO存在明显的负相关.在所有大型底栖动物中,甲壳动物26种,多毛类环节动物24种,软体动物23种,棘皮动物8种,其他类群动物6种(鱼类5种,尾索动物1种).该海域的优势种为多丝独毛虫和短叶索沙蚕.将CLUSTER结果以43%的群落结构相似性来划分,组内的相似性为50%~60%.该海域大型底栖动物的平均丰度为854个/m2,平均生物量为134.73 g/m2,与研究海域附近的历史研究结果相比较,该海域的平均丰度和生物量仍处于较高的水平.     

Comparative study on the fauna composition of intertidal invertebrates between natural and artificial substrata in the northeastern coast of Jeju Island

This study was carried out to learn about differences in the sessile macrobenthic fauna communities between the artificial and natural habitats. There were some differences in terms of species composition and dominant species and community structure between two habitat types. The dominant species include Pollicipes mitella and Granuilittorina exigua in natural rocky intertidal zones; Monodonta labio confusa, Ligia exotica, Tetraclita japonica in the artificial rocky intertidal zones. Among all the species, L. exotica and T. japonica occurred only in the artificial rocky intertidal zone. The results of cluster analysis and nMDS analysis showed a distinct difference in community structure between artificial and natural rocky intertidal zones. The fauna in the natural rocky intertidal zones were similar to each other and the fauna in the artificial rocky intertidal zones were divided depending on the slope of the substratum. In the case of a sloping tetrapod, M. labio confusa and P. mitella were dominant, but at the vertical artificial seawall, Cellana nigrolineata, L. exotica T. japonica were dominant. The analysis of the species presented in natural and artificial rocky intertidal areas showed the exclusive presence of 10 species on natural rocks and 12 species on artificial rocks. The species in the natural rocky intertidal area included mobile gastropods and cnidarians (i.e. rock anemones), and the species in the artificial rocky intertidal area mostly included non-mobile attached animals. The artificial novel structure seems to contribute to increasing the heterogeneity of habitats for marine invertebrate species and an increase the species diversity in rocky coastal areas.     

乐清湾大型底栖动物群落及其与环境因子之间的关系

为了解和掌握乐清湾大型底栖动物群落及其与环境因子的相关性,于2015年1月、3月、5月、8月和10月开展了5个航次的大型底栖动物和环境因子的调查。结果表明,1月、3月和5月3个月份的群落结构和空间分布均较为接近,优势群落的主要特征种为薄云母蛤和不倒翁虫;8月和10月的优势群落不明显,主要群落特征种有不倒翁虫、双形拟单指虫、寡鳃齿吻沙蚕、小头虫、中蚓虫属一种等。丰度/生物量曲线(ABC曲线)分析表明5月的群落结构较稳定,1月、3月、8月和10月均处于不同程度的扰动状态,其中8月的受扰动程度最大。BVSTEP分析表明亚硝酸盐是乐清湾大型底栖动物群落各月份差异的主要环境驱动因子,盐度和氨氮分别是影响乐清湾3月和5月大型底栖动物群落结构的主要环境因子。典范对应分析(CCA)表明水温对乐清湾大型底栖群落特征种时空分布影响最大,其次是盐度、溶解氧、含氮营养盐和pH等。通过CCA排序图发现大部分群落特征种分布在含氮营养盐较低的水域,表明高浓度的含氮营养盐已经对乐清湾的大型底栖动物产生了负面影响。     

山东长岛潮间带大型底栖动物生态特征研究

2015年11月(秋季)、2016年2月(冬季)、2016年6月(春季)、2016年9月(夏季)大潮期间对山东长岛列岛中南长山岛和北长山岛的3个不同底质类型的潮间带采样点进行了系统的大型底栖动物调查,研究了其数量特征和季节变化,包括群落种类组成、密度和生物量、优势种、物种多样性等,结果表明:本次调查共鉴定出大型底栖动物隶属7个门,共81种。其中环节动物门多毛类动物25种,软体动物门25种,节肢动物门22种,棘皮动物门6种,其他类群共3种(包括纽虫、腔肠动物和鱼类各1种)。潮间带大型底栖动物平均密度为118.97 ind./m~2,软体动物和甲壳动物占据了研究区域底栖群落的密度主要成分,两者分别占比为39.58%和36.68%。各采样点平均密度为北长山岛(BCS)(141.17 ind./m~2)避险角(NCS1)(122.33ind./m~2)明珠广场东(NCS2)(93.42ind./m~2);各采样点平均生物量为128.03g/m~2,软体动物和甲壳动物占据了研究区域底栖群落的生物量主要成分,两者分别占比为58.56%和35.32%。平均生物量北长山岛(BCS)(234.95g/m~2)避险角(NCS1)(99.99g/m~2)明珠广场东(NCS2)(49.13g/m~2)。从季节来看,各站点的物种数、平均密度和生物量季节变化的规律性不明显,总体为秋、春季较多,夏、冬季较少。3个站点物种多样性指数(H′)、丰富度指数(D)和均匀度指数(J′)平均值分别为2.30、1.99和0.79,各潮区中,均为中潮带低潮带高潮带,各季节变化不明显。优势种受底质类型和季节变化的影响明显,没有在各站点和季节均为优势种的种类,主要的优势种有平背蜞(Gaetice depressus)、绒螯近方蟹(Hemigrapsus penicillatus)、短滨螺(Littorina brevicula)等。与以往历史资料相比,长岛潮间带生物出现了一定变化,种类和数量减少,与烟台近岸潮间带大型底栖动物种类组成差别很大。     

Trophic structure and production in the macrobenthos of a temperate Australian estuary

The soft-bottom fauna of Gunnamatta Bay, Port Hacking, N.S.W., was sampled to estimate the abundance, trophic structure and net production of the macrobenthos. The samples were classified into groups from spatially distinct strata, the groups differing in species abundance and diversity and in the identity of the dominant species. The composition and community structure of the fauna was generally similar to that in other sheltered embayments on the southeastern coast of Australia. Estimated production values were low for an estuary, ranging from 10 kJ m^?2 year^?1 in an intertidal stratum of well sorted sand to 210–450 kJ m^?2 year^?1 in shallow subtidal strata with an admixture of silt. Estimated net production over all strata was 160 kJ m^?2 year^?1. Polychaetes dominated production, providing 45% of the total; crustaceans provided 20%, and molluscs and echinoderms 12% each. Detritus-feeders were dominant in most strata: the bivalve Theora fragilis, the polychaetes Australonereis ehlersi and Notomastus torquatus, and the urchin Echinocardium cordatum were major contributors to overall biomass and to net production. Substantial biomass and net production was also contributed by the suspension-feeding bivalve Corbula vicaria and a carnivorous crab Ceratoplax cf. glaberrimus. Overall, detritus-feeders contributed 61% of estimated net production, carnivores 32% and suspension-feeders 7%. Production estimated for carnivores was higher than could be supported by the non-carnivore populations, suggesting that some of the species considered to be carnivores have additional feeding modes. Excessive harvesting of bivalves by humans is suggested as the main reason for the low biomass of, and therefore low production by, suspension feeders.     

Macrobenthic communities of the Vellar Estuary in the Bay of Bengal in Tamil-Nadu in South India

The macrobenthic fauna and communities of the Vellar Estuary located at the southeast cost of India (11°30′ N, 79°45′ E) and the adjacent marine and river habitats are described on the basis of original data (70 samples over 10 transects). The fauna consists of 115 macrobenthic species and 79 species in estuarine habitats. We described 14 types of macrobenthic communities with different compositions of the dominant species. The leading ecological factors of the distribution of the communities are the salinity, depth, and bottom type. The Vellar estuary consists of two longitudinal zones of macrobenthos. The polyhalinic area is populated by the marine species, but it is related not to a salinity decrease but to the protection from waves and silt on the bottom in this area. The polyhalinic communities are most abundant in terms of the biomass and species richness. The mesohalinic area is inhabited by brackish water species and communities with low abundance. The sublittoral estuarine area is dominated by filter-feeders—the bivalves Crassostrea madrasensis, Meretrix casta, Modiolus metcalfei, and Scapharca inaequivalves—and the littoral zone is dominated by the gastropods Cerithidea cingulata, some crabs, and polychaetes. The ecosystem function of the Vellar estuary can be defined as a filter for the fine organic particles transported by the river.     

Influence of Environmental Variables on the Distribution of Macrobenthos in the Han River Estuary, Korea

We compared environmental effects on the macrobenthic community of the Han River Estuary in summer, when freshwater input from the Han River increased, and in spring, when freshwater input decreased. Field samples were taken from the upper region of the Shingok reservoir to the southern area of Ganghwado at 18 sampling sites after rainy (August 2006) and dry (March 2007) seasons. Macrobenthic fauna were collected using a Van Veen Grab (0.025 m² and 0.1 m²) and environmental factors were measured simultaneously. Dominant species of macrobenthic fauna and the macrobenthic community were divided into two areas, the area of the Han River with no salinity (< 0.1 psu) and the southern part of Ganghwado with salinity (> 20 psu). The dominant species Byblis japonicus appeared at Junruri in the dry season. The distributions of two polychaetes, Hediste japonica and Nephtys caeca, were divided into the lower and upper areas of the Singok submerged weir. BIO-ENV (the matching of biotic to environmental patterns) analysis revealed that salinity was the most important factor affecting macrobenthic communities in the Han River Estuary, with other factors such as sediment grain size, bottom dissolved oxygen, and total organic carbon of sediment being secondary.     

Gorgan Bay:a microcosm for study on macrobenthos species-environment relationships in the southeastern Caspian Sea

The relationship between spatial patterns of macrobenthos community characteristics and environmental conditions(salinity, temperature, dissolved oxygen, organic matter content, sand, silt and clay) was investigated throughout the Gorgan Bay in June 2010. Principal components analysis(PCA) based on environmental data separated eastern and western stations. The maximum(4500 ind./m²) and minimum(411 ind./m²) densities were observed at Stas 1 and 6, respectively. Polychaeta was the major group and Streblospio gynobranchiata was dominant species in the bay. According to Distance Based Linear Models results, macrofaunal total density was correlated with silt percentage and salinity and these two factors explaining 64% of the variability while macrofaunal community structure just correlated with salinity(22% total variation). In general, western part of the bay showed the highest number of species and biodiversity while, the highest density was found at Sta. 1 and in the middle part of the bay. Furthermore, relationship between diversity indices and macrobenthic species with measured factors is also discussed. Our results confirm the effect of salinity as an important factor on distribution of macrobenthic fauna in south Caspian brackish waters.     

胶州湾春季大型底栖生物多样性及群落结构特征

基于2018年5月胶州湾海域共计20个站位所获取的生物及环境数据，研究胶州湾大型底栖生物多样性以及群落结构特征。调查海域共发现大型底栖生物208种，大型底栖生物总平均丰度和生物量分别为2 654.38 ind./m²和1 024.512 2 g/m²，大型底栖生物物种数、丰度和生物量均呈现北部高、南部低的分布趋势；调查海域优势种以多毛类为主，I_RI值最高的物种为菲律宾蛤仔Ruditapes philippinarum；香农维纳指数与ABC曲线均表明调查海域底栖生态环境整体清洁；在40%相似性水平上，可将调查海域大型底栖生物划分为4个群落；底温和底盐影响调查海域大型底栖生物丰度分布。     

胶州湾大型底栖动物的丰度、生物量和生产量研究

为了研究胶州湾大型底栖动物的丰度、生物量和生产量,于2002年3月、6月、8月和12月,在胶州湾北部软底区、大沽河口、黄岛养殖区及养殖区邻域选取4个站位进行采样,对大型底栖动物进行了定量研究。共采到大型底栖动物138种,总平均丰度、平均生物量(湿质量)和年生产量(有机碳)分别为1 719个/m2,27 g/m2,2.2 g/(m2.a),初步估算,胶州湾大型底栖动物的总次级生产量为2.8万t/a。与渤海和南黄海大型底栖动物的丰度和生物量比较,丰度和生物量均低于这两个海域,但是胶州湾大型底栖动物的总次级生产量高于渤海。本研究对于了解胶州湾大型底栖动物现状及湾内养殖对大型底栖动物的影响具有重要的意义。     

杭州湾大型底栖动物季节分布及环境相关性分析

对杭州湾大型底栖动物进行了夏(2006年7-8月)、冬(2006年12月-2007年1月)、春(2007年4月)、秋(2007年10-11月)季调查,分析了大型底栖动物的季节分布及其与环境的关系。结果表明:本海域四季共鉴定出113种大型底栖动物,多毛类、软体动物和甲壳类是该海域大型底栖动物的主要类群;主要优势种为双鳃内卷齿蚕、西方似蛰虫和多鳃卷吻沙蚕等多毛类。方差分析表明,除甲壳类丰度外,杭州湾大型底栖动物群落参数无显著季节差异,群落分析表明,杭州湾群落特点比较单一,未分辨出显著差异的群落;丰度/生物量比较法分析表明,秋季杭州湾海域受到了污染的扰动;典范对应分析表明,多毛类主要受物理因子(如温度、盐度、水深等)影响,而软体动物、甲壳类和其他类动物主要受生物化学因子(如叶绿素、总有机碳、浮游生物等)影响。从典范对应分析的结果,可以推断出杭州湾大型底栖动物受自然环境特征的影响较人类活动的影响大。     

澄迈马袅湾网箱养殖区及周边海域大型底栖生物调查

本研究分别于2019年7月(夏季)和12月(冬季)对澄迈马袅湾网箱养殖区及周边海域的大型底栖生物进行调查。结果表明,两次调查共采集鉴定到55种生物,其中夏季31种,主要分布在邻近区;冬季33种,主要分布在养殖区和邻近区。夏季航次,底栖生物在养殖区、邻近区和对照区的栖息密度分别为30.55 ind/m²、40.74 ind/m²和29.63 ind/m²,生物量分别为14.36 g/m²、6.28 g/m²、6.58 g/m²;冬季航次,三个海区底栖生物栖息密度分别为55.56 ind/m²、46.67 ind/m²和27.78 ind/m²,生物量分别为7.29 g/m²、6.20 g/m²和0.22 g/m²。夏季优势种主要为绒毛细足瓷蟹和纽虫;冬季优势种主要为豆形短眼蟹、梳鳃虫、纽虫和哈氏美人虾。夏季养殖区、邻近区和对照区大型底栖生物多样性指数(H′)分别为1.40、2.06和1.46;冬季分别为2.16、1.59和0.94。综上分析,网箱养殖产生的残饵和鱼类粪便,对养殖区及邻近海域底栖生物分布产生一定影响,且其群落结构与区域和季节存在较强相关性。     

杭州湾和三门湾拖网大型底栖动物群落组成和多样性研究

本研究根据2017年8月和2018年8月在东海北部杭州湾和三门湾大型底栖动物的拖网采集资料,对这两个海域的大型底栖动物群落结构和多样性进行了研究。杭州湾的10个站位和三门湾的6个站位共获得61种大型底栖动物,其中甲壳动物最多(22种),其次是鱼类(20种)、软体动物(14种)、棘皮动物(2种)、刺胞动物(2种),多毛类最少(1种)。杭州湾大型底栖动物优势种出现3种,三门湾出现8种,脊尾白虾Palaemoncarinicauda是两个海域唯一的共同优势种。相比2017年,杭州湾和三门湾在2018年鱼类的种类和数量都有明显的增长。根据大型底栖群落的多样性指数,杭州湾不同区域的生物多样性差异较大,而三门湾则较为平均。对杭州湾和三门湾大型底栖动物进行聚类分析和nMDS排序分析,结果表明,杭州湾不同站位大型底栖动物相似度低,Q8、Q9、Q10站位群落结构与三门湾的站位更为接近,三门湾的底栖群落在2017年和2018年存在较大差异。杭州湾和三门湾大型底栖动物群落的变化可能与环境因子、人类活动、厄尔尼诺事件等的变化相关。     

Prey versus substrate as determinants of habitat choice in a feeding shorebird

Many shorebirds on their non-breeding grounds feed on macrobenthic fauna which become available at low tide in coastal intertidal flats. The Eastern Curlew Numenius madagascariensis in Moreton Bay Australia, varies greatly in density among different tidal flats. This study asks: how important is the abundance of intertidal prey as a predictor of this variation? We quantified feeding curlews’ diet across 12 sites (different tidal flats, each re-visited at least eight times), through 970 focal observations. We also estimated the abundance of total macrobenthic fauna, potential prey taxa and crustacean prey on each tidal flat; measured as the number of individuals and a relative biomass index per unit substrate surface area obtained from substrate core samples. We estimated curlew density at each site using low-tide surveys from every site visit.