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    6 views176 pages

    Change in Marine Communities

    Uploaded by

    neta
    primer v6 guide

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    © All Rights Reserved
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    CHANGE IN MARINE COMMUNITIES: An Approach to Statistical Analysis and Interpretation 2nd Edition K. R. Clarke & R. M. Warwick Cum Plymouth Marine Laboratory, UK -E Lid Pablished 2001, by PRIMER-E Lid Plymouth Marin Laboratory Prospect Place West Hoe Plymouth PLI 3DH United Kingdom Business OFice: 6 Hedingham Gardens Roborough Plymouth PL6 7OX United Kingdom ios K Raber Cee MSOPRD_Rayind Gy MA Reprimed 957 Sond etn 2001 ‘Clarke, KR, Warwil, ML, 2001 ‘Change it marine communities: an approach to statistical analysis and interpretation, Dad odeon. PRIMERCE: Plymouth Copyright 2001 PRIME E Li al igh esd Cover phos: Steve Smith Unerty f New England Armidale, NSW, Aas, Pl Somerfield (Pymoth Marine Lert, UN), and hey Remon NDA ligt, No Zesand CONTENTS INTRODUCTION CHAPTER 1 CHAPTER 2 CHAPTERS cuaprer ‘CHAPTERS (CHAPTER 6 CHAPTER 7 (CHAPTERS CHAPTERS CHAPTER 10 (CHAPTER 11 carrer 12 (CHAPTER 13 CHAPTER 14 CHAPTER 15 CHAPTER 16 cHaPreR 17 APPENDIX APPENDIX APPENDIX ‘A framework for studying changes in community structure “Measures of sinilariy of species abundance biomass between samples, Hierarchical clustering Ondination of samples by Principal Components Analysis (PCA) Ondination of samples by Muli-Dimensional Scaling (MDS) Testing for differences between groups of samples Species analyses Diversity measures, dominance curves and other graphical analyses Transformations Species removal and ageregaton Linking community analyses wo environmental variables Causality: community experiment in the fel and laboratory Data requirements for biological effects studies: which components and attributes of the marine biota to examine? Relative sensitivities and merits of univariate, graphical/distributional and ‘multivariate techniques. Multivariate measures of community sess Further comparison of multivariate patterns Biodiversity measures based on relatedness of species Index of example data Principal literature sources and further reading References cited INTRODUCTION Parpose ‘This manual accompanies the computer sofware package PRIMER (Plymouth Routines in Maltvariate Fcologieal Research), obtainable from PRIMER-E Ld, Plymouth see wow primer som). le scope i the analysis of data arising in community eslogy and ‘ervionmertal scence which fs mula ia character {onany species, multiple environmental variables) and its intended for use by ecologists with no more han a minimal background in statistics. AS sch, thi methods ‘manual complements the PRIMER user minal, by ving the background to the statistical whic employed bythe analysis programs (Table?) t level of detail hich shot allow the eicogist 10 lunderstand the Outpt from the programs, beable 0 Aesrie the results ina non-technical way to others tnd have confidence thatthe right methods ae being sed forthe right problem. “This may seem a tll order, in an area of statistics (primarily multivariate anal) which septation 4 esotrie and mathematically complex! However, ‘whist itis tre thatthe computational details of some Of the core techniques described here (For sxample, on-met multidimensional sealing) are decid n= trivial, we maintain that all of the methods at have Tale Chap tehsil Inerbng see PRIMER einer are ever Rowtnes ‘Capes ‘Sinan a ‘CLUSTER 3 Pca an os “anos ANOSTI 6 sire 1 DiveRse an CASWELL, Goomertc & Dominance Pots '§ Tranjerm 9 Atarewase 0 ‘BIO-ENY, Draiuman Plot " IMYDISP, RELATE 5 BUSTEP, STAGE 6 peer creeper joaemesahyae eins Introdeton ree -| ‘ben adopted or developed within PRIMER are so conceptually seaightorward a to be ameneble to ‘simple explanation and transparent interpretation In fact, the adoption of non-parametric and permutation approaches for display and testing of multivariate ata requires, paradoxically, lower level of statisti Sophistication onthe pr ofthe usr tan ds satis- factory exposition of standard (parametric) hypothesis testing inthe wnivoriae case, ‘The principal am ofthis manus is therfore to dsibe 4 coherent strategy for the interprettion of dita on ‘community strutore, namely values of abundance, biomass, % cover presencefabecace ee, fora et oF species and one oF more replicate samples shen 8) ata numberof sites atone time spatial analysis), 1b) atthe same site at a number of times (temporal analysis); ©) for @ community subject to different uncontrolled ‘or manipulative "weatment'; or some combination ofthese, These speceshy-samplesarays are typically large, and patems in community ste ate often not ey pparent fom sinpe inspection of the dts ‘Statistical tals therefore cea around redoing the compl iy ofthese matrices, sll by some graphical pres tation of the biological relationships between the samples. This is followed by statisti! testing 19 ‘eri and characterise change in community sacture Intime or space and ela these to changing environ ‘mental or experimental conditions, Material covered 1 should be made clear atthe outset that the te "Change ia Marine Communities” doesnot in ny ‘vay reflect a restrition inthe scope ofthe techniques in the PRIMER package to the marine environment, ‘The fist eiton of this manual war intended primarily for a marin audience and, given that ts examples are sill drava entirely from marine contexts, would be disingenuous to change the tile ow toa more general fone. However, it wll be self-evident to the reader ‘that there is very litle in he following pages tat ‘exclusively marin, Indeed, the PRIMER package i ‘now not only used world-wide forall types of maine ‘community surveys and experiments, of bethi ana, alge, corals, plankton, fis, diet stadies ete, but is ineresingly found in freshwater, tenes and palacontlogy contexts, and sometimes solely in mul ‘arate studies of physico-chemical characteristics, ‘As result ofthe authors’ own esearch interests and the widespread use of commnity data in polition ‘monitoring. amor isto the manual ithe Biological fet of contains but, again, mos ofthe methods fre much more genealy applicable. Thisis reflected ina range of more fundamental ecological studies song the eal datasets exemplified here ‘The leramre contains a large array of sophisticated staistical techniques for handling species-b- samples atc, ranging rm tee redacton wo spe dere sy. indies, through curvilinear or distributional representations of richness, dominance, evenness et, ‘2 plethoa of multivariate approaches involving lasing or ordination methods. This manual does hot attempt i give an overview ofall the options, oF ven the majority of them. Instead it present satay ‘which has evolved over several years within the Community BcologyBiodivesity group a Pimouth Marine Laburatory (PML), and which basa proven teak eco in interpretation of a wide ange of marine community cata: se, fr example, papers listed under {Clarke or Wersick in Appendix 3 (which have attained fouefigure tal ctions i SC journals). Tho analyses in displays thse papers, andi his manual almost raw up the wide ange of routines available in ‘he PRIMER package hough in many cases amnotations ‘ten plots hve boo Further elite by simple ping ino graphics pogems suchas Microsoft Pwverpiat). Note also that, whist other software packages wi not encompass this specific combination of routines, everal of the individual techniques can be found elsewhere. For example, the core clustering and ‘ordination methods described here ae available in ‘Several munstream statistical packages (SAS, -Phs, Syst Statgaphies ete), and more speed stat- isc’ programs (CANOCO, PATN, PCORD, the Cornell Eedogy programs, et.) tackle essentially similar probes. though usally employing diferent techniques and a diferent strategy ttalning wokshops funded jointly by FAO, UNEP and UNESCOMIOC, and a series of commercially-un PRIMER courses at Plymouth and venues outside the UK. The acvocacyof these techniques ths springs not nly rem vegular use and development within PML's Community Esology/Biodiversty group but leo fo valiablefecdback fom a seis of workshops in which rate data analyses were central ‘Throughout the manual extensive use is made of data sets rom the pulse iterate to illustrate the et niques. Appendix 1 gives the cial erature sour foreach ofthese 25 o so data Sets and an index tall the pages om which they are analysed, Each dataset is allocated singe leer designation an, to avold confusion, refered fin the text ofthe mal by tha let, placed in curly brackets (eg. 4) = Amoco Cadiz sil spi, macrofauns; (8) ~ Brisol Chanve, 200- ‘ankton: (C7 = Celie Se, zooplankton ete. Literatare citation ‘This 2nd edition ofthe manual follows the Ist ition elosely in espect ofthe Fist 15 chapters, though minor revisions have been made throughout” Chapters 16 tnd 17 ae entirely new. Appendix? lists some back- ground papers appropriate o each chapter, incling the souree of specific analyses, ard fll isting of referenees cited isin Appendix 3 ‘Whi the manual is peninely collaboratively authored, forthe purposes of drestng queries on specie topics ‘tis broadly tre that the rst ather (KR) Bears the responsibility forthe chapters on satstical methods (077, 9,11) and the second author RMW) i mainly responsible forthe chapters on nt pretation (10, 12 14), the responsibility for Chapters & and 1S being shared more or less equally. Chapters 16 and 17 were ‘writen by KRC, drawing onthe rests of joint popes in various authorship combination by KRC, RMW and Paul Somerfield (also of the Plymouth Marine aboraton). Since this manual is ot accessible within the published literature, referal 0 the methods it destibes would properly be by citing the primary person which itis based: these ae indicated in the text and Appendix 2. Alternatively, comprehensive Aiscussion ofthe philosophy (and many of the details) ofthe molvviat and univariate approaches abvocted fan be found in Clarke (1993, 1599) and Warwick (1993, respectively, withthe newer methods in his tition best summarised in Clavke and Warwick (19983), Somerfield and Clarke (1995) and Warwick nd Clark (2001). Acknowledgements ‘We are gratefil to a large numberof individuals and Insitutions for thee help and support ~ pease se the etal ist tthe end ofthe mani KR Clarke RM Warwick 2x01 Chapt age CHAPTER 1: A FRAMEWORK FOR STUDYING CHANGES IN COMMUNITY STRUCTURE ‘The purpose of this ofeing chapters twofold 8) to introduce some ofthe datasets which are used ‘extensively as illsrations of techniques, trough- ‘ut the man 1) to outline a famework forthe various posible sages in a communi analysis. Examples are given of some core elements of the recommended approaches, foreshadowing the analyses ‘explained in detail ner an referring forward othe relevant chapters. Though, a this stage, the details fre Tikely to remain rysifpng, the intention f that ‘his opening chaper sould give the reader some fee! Tor where the various zchniqus are leading and how they slot topeter. AS such, ii intended to serve both as an introduction and a summary Stages tis convenient ctezoris possible analyses bray int four main stages 1) Representing communities by fp description ofthe relationships between the tain the various Samples. This i bought of a8 pure description, rather than expanaton etext, ad the emphasis iS on reducing the sompleity ofthe maltivariate information in typical spcie/simples matrices, 2 ‘obtain some form af low-dimensional piste of how the biological amples ntereate, 2) Discriminating steveondtions onthe basis of hci biotic composition. ‘The paradigm her i that of the hypothesis test, examining Whether there are “proven” community differences beoveen groups of samples identified prior for example demon- stating diferences between contol and putatively Impacted sites, esablishing beforlaftr impact ferences ata singe site ee 3) Determining level of “sess” 0 disturbance, by tempting to const biological measures from the Community data which ae indiestive of disturbed ‘conditions. These nay be absolute measures this ‘sbeered scr! fue is ndtve of polaion”) ‘or relative eritria under impact, this coocient “rhe commun waned oh he re ment ose oceans ang ae ‘wear imply sna orci pen spoon cmp tapes fas is expected to doors level") Noe the con fe previous stage, however, which i estetd 1 demons ones betwee grup of samples, not asrbing Airetionalty to the change (eg. deleterious sone sequence). 4) Linking to environmental variables so exasiving ‘sues of enusaiy of any changes. Having allowed the bilogies! information to “lls onn ston”, any associated physical of chemical” variables ‘matched the same set of samples can be examined for thei own structure and its relation othe Bote patem (is “explanatory power"), The extent 10 ‘hich identified environmental differences are actually eamsal to observed community changes can only relly be determined by: manipulative experiments, either inthe field or though laboratory Frmesocosm tues, ‘Techniques ‘The spread of methods for exacting workable repre ‘mations and summaries ofthe biological data can be srouped into tne categories, 1) Univariate methods collapse te ful sto species ‘counts fora sample into single coefficient, for example a species diversiy index. This might be Some measore of the numbers of different species fora fixed pumber of individuals (species richness) forthe exten o which the community counts are dominate ty a sal numberof species (dominance Fevenness index), of some combination of these Also included ar lovers indices which measre the degree to which species or organs in a sample are taxonomically or phylogenetically related to eachother. Clea, the prior selection ‘of asingl taxon as an indicator species, amenable to specifi inferences about its response toa partic- lar environmental gradient, alo gives rise fo.@ ‘ivr nasi. 2) Distributional techniques, sso termed graphical ‘or cunilinear plot (when they are not strictly Aistutona, “are a elass of methods “which summarise the set of species counts for single sample by a curve or histogram. One example ise dominance curves (Lambsad eo 1983), bic rank the species in decreasing order of abundance, ‘convert the values percentage abundance relative page 1-2 to the tot number of individuals inthe sample, fand plot the cumulated percentages against the Specs rank, This, andthe analogous plot based (of species biomass, are superimposed to define ABC (abundance-biomass comparison) curves ‘Warwick, 986), which have proved a sof com ‘retin investigating disturbance eflets. Another example is the species abundance distribution Gomatines termed the dsrbution of individuals lamongstspriet) is which the species ae cates ‘red ino geometrically sealed abundance classes land a histogram plated ofthe number of species falling in each abundance range (eg. Gray and Pearson, 1983), tis then argued again from emp inal evidence, hat there are cern characteristic changes in fis distribution associate with comm ‘nity disturbance, ‘Sach distribatona techniques rel the constraint inthe prevous category tht the summary from cach sample shouldbe a singe variable; here the emphasis is more on diversity curves than single ‘iversity ines, bu note hat both these catagories share the propery tha comparisons between samp les are not Based on particular species identities: ‘to samples can have exactly the same diversity or Alistbutional structure without possessing a single species in exmmen, 5) Multivariate methods ar characterised by the Fst that they be their comparisons of two (or more) samples on he extent to wich these samples share particular species, at comparable levels of abund- ance. Either explicitly or implicitly all molt techniques se founded on such similarity ceffic- ents, clelaed beeen every pair of samples There then facilitate a classical oe clustering (these terms are interchangable) of samples into soups which re mutally similar, or an odinaion loin which or example, the sample re “mapped” (swallyin two oF tre dimensions in sucha way thatthe distances between pairs of samples veteet their relative dissimilarity of species compsition Techwigues described in detail inthis mans! are 9 Imethod of erarchica agglomerative clustering (ea. Eveit, 1980), in which samples ae suscessive- Iy fused into larger groups, av the erterion forthe similar Tevel defining group membership i relaxed, tnd evo ordination techniques: principal components analysis (PCA, eg. Chatfield and Collins, 180) and tnom-meric mut-dimensonal sealing (NNIDS,wsuly Shortened to MDS, Kruskal and Wish, 1978) For each broad category of analysis, the techniques appropiate to cach stage are now diseusie, and pointers piven othe relevant chapters. UNIVARIATE TECHNIQUES For diversity indices and other singlevarsble extractions from the data marx, standard satis ‘methods are usually applicable and the wader is refered to one of the many excellent general statisti texts (eg Sokal and Rohlf, 1981) The equisite techniques fo each tage ae summarised in Table 1.1. For example, when samples have the structure of @ numberof replicates taken at each of 3 umber of sites (or times, or conditions), computing the means and 95% confidence intervals gies at appropriate represemtation of the Shannon diversity (Gap) at each ste, with discrimination berwoen ster being demonstrated by one-way analysis of variance (ANOVA), which sa test ofthe mull hyptteis that there ae no differences in mean diversity between ale Unt teenie. Sammars foro es Ua pe Does ies (Or) adr her nentCh 7) 1 Reps eon nd 95% conden mri oh steondton Ch, 9.7) 2) Distniting se ons far ANOY sites‘condiions Cnorey eeapae es 2) Deeminige Byrne hr! ear te ( 1,19) regina “pec po041) Srarlees| —Unimay aden mcs al meete ln operant Lat of oom tne 2 Lakin Aarne O41): focal ince Ch 12 sites, Linking tthe environment is ten als relt- ily staihtfrward, parca i th environmental ‘arabes can be condense int one (osm number «key summary statistics. Simple or mute regres fan of Shannon diversity asthe dependent variable, Against the envionment descriptors as independent ‘ariabls, i then technically feasible, though rarely SE infrmtine n act, en the needed rate ofthe information ut For impact studies, much hasbeen writen about the ect of potion or disturbance on diversity mesures hist the eesponse is not necessarily undiectionl (ander the hypothesis of Huston, 1979, diversity is expected to rise at intermediate disturbance levels tefore its strong decline. with gross disturbance), there is sense in which determining stress levels is pps, hough elon to historia diversity paterss for particular environmental gradients. Similarly, teplrical evidence may exist that particlar indiestor texa (@e. Capitli) change in’ abundance along spe olluton gradients eof oginc enrichment), [Nate though that, nlite the diversity measures con- structed fom abundances across species, averaged in some was indicator species levels or the number of fecies ina sample (S) may not intl ste the ‘sumptions necessiy for clasizal statistical analysis For, the nomality and constant variance conditions can usually be produced by tansfomation of the variable (ele 8). However, for most individu species abundance aos these of samples i key XE be a very poorl-behaved variable, statistically syeaking. Typically, species wil be absent from any ofthe samples and, when tis present, the counts ‘ae often highly variable, with an abundance probab- ity disebuton which is eavly rightshewed! Thus, ferall ba the most common individual species trans. Fermation is no real help and parametric statistical analyses canna be appli tothe counts, in any or, In any ease, iis not Valid to “snoop” in a large data natrix, of typically 100-250 ta, for one oF more “interesting” species to analyse by univariate techn- ‘gues (any indeator or heystone species selection must Anda ajc ema ihrem, wih en "at r,t in he ly fe nmani, ha sof pees a dated ‘iron pea Plon pratt a nh cee ‘a ie Howat lc! trator foci tet ‘Chaser mechani of ern, warty ey ‘eect. "Ta ear cou mae teed onerliered contin wit igh sof ‘rac cing mae oon aping prams mig ‘eweoreal nto Chapter 1 meets be done a prior. Such arguments lead to the tenets underlying this mana {community dita re usually highly mivarite (large numbers of species, cach subject 10 high Statistical mole) and need tobe analysed en masse in order (0 ect the important biological signal nd it elation to he environment ‘standard paramedic modelling stotaly inva ‘Thus, throughout litle emphasis given to represent ing communities by univariate measures though some Aetnitons of iedices can be found atthe sat of Chapter 8, some bref remarks on hypatheris testing (ANOVA) atthe start of Chaper 6,8 discussion of transfematons (» approximate aot ac constant sariance) at the tart of Chapter 9, an example given ofa univariate regression between biota and environ ment in Chapter 1, and a moee extensive discussion of sampling properties of diversity indices and bi diversity measur based on taxonomic relatedness, ‘makes up Chapt 17. Finally, Chapter gives a series of detalled comparisons of tnivarate with Aistrbutonal and multivariate techniques in onde to ‘sauge their elative sensitivities and merits na range of practical studies, EXAMPLE: Fvierfjord macrofauna ‘The first exampye is fom the JOCIGEEP practical workshop on bisogical effects of plltants (Bayne tal 1988), hel atthe University of Oslo, August 1986, Thisatenyeed o contrast a range of biochem, > 1) ant the prevalence of zeros Here, as slsewhere, even an undesirable reduction to the 30 “most important” species (se Chapter 2) leaves more Chapter ig 1. Peer Nene IF. Bc onion sctihane CURED Dua Wap teamed Jeena yt ary ‘Pig. 12. Poeerd mcrfaana fF). Maas ad 9% fice trator Sarr es om ar ‘plea aeach afar) than 50% of the matrix consisting of zeros. Standanh multivariate normal analyses (eg: Maria eal 1979) ofthese cots are ceri ruled ou; they require both that the number of species (variables) be small in relation to he numberof samples, and that the abund- !nce or binmass values are transformable 1 approx mate norway nether is possible. As discussed above, one easy route to simplification ofthis “high-dimensional” complexity i to reduce tach column ofthe matrix (each sample) to single, Tite 12, Pernt micrfaun Ph Arion ones ics at ny ore 0 secs 4 amps op era fs 4) ebmdoee nabs or (ne oma tg po —__ Semper oa ALAD AS AG Ba Abundance Corioniualions 0 0 00 0000 Halenpusp = 0 0 0 1 D000 Onchnsoms = 0 0 OD DOOD Phacalonsvonbs 0-0 0-1 9010 a Holohwodee = 0 000 «9000 Nemerina nde 12-6 «8 6 9 61? Pobeoua mi = 3 000 0010 Amocnariaa 1 1 10 9000 Anphietesgumed 0 0 0.0 4 0.00 Anpharcie «0 000 1000 Anatidesgrom. «0 0 OT 1000 acter 0000 0000 Biomass Conanturlinds 0 000 0000 HHatenpmssp = «0 0 0 0 000 JOncineroma = «0 0 0 0 0 OOO Phascolonsrombi 0 0:0 6 0 020 a a Hoohuroeo = «0 9-0 0 0 O00 Nemertina det 140390 «1S TOT Popcerm inde «= 290-0 0 0 0 OO JAmscna rotate 4 14234-00800 JAmphces sumer -0-0«0:«0 «0 00 Amphoe «0:0 0 0 0 OOO JAnandesgroen. «=«0:«0:«0 FT «HO OO JAnawdse 0 OD OOOO univariate descriptin. Fig. 12 shows the results of computing the Shanton diversity (H, see Chaper 8) of each sample’, an ploting for each site the mean diversity and its 95% confidence interval, based ona pooled estate oftariance acrost al sites from the ‘ANOVA table, Chapter (An analysis ofthe type ‘outlined in Chapter shows that prior ransformation (of HT is not requir it already has approximately ‘constant variance acres the ites, a necesary pees ie for standard ANOVA). The most obvious feture of Fig. 12 isthe wlatively higher diveesity atthe conto” lation, 4 1 ng the PRIMER DIERSE rate Stazes ‘ABC dominance) ares (Ch) 1 Representing Cares for och 2) Dissininating ANOVA ona Sends ANOSIM tex (Ch6) on “dsc erween eer pa of cues 3) Determining Biomass ere shops Blow stessleves mbt curve wer dtbance 4) Liking to Dia exp for ai DISTRIBUTIONAL TECHNIQUES ‘A les condensed frm of summary of each sample is ‘ered bythe dsetional graphical methods utined Forte four stages in Table 13. Representation is by curves or histograms (Chapter 8) either ploted foreach replicate sample spaately or fer pooled data witha sites or conditions. The former Permits visu judgement ofthe sampling raciation Inthe curves and, as with diversity indices, plication require to dscriminae sie, ic ts thew yp ‘ss tat two or more sites (conditions et) have the ‘ame curvilinear structure. The easiest aproech to ‘esting then to summarise each eeplizate eve by 8 single statistic and apply ANOVA as befoe for the ‘ABC mathod, mentioned eat, te I staisiChapter 8) isa convenient measre ofthe extent to which the biomass curve “dominates” the abundance carve, or ce-vesa. This is efective in practice though, in theory simply amounts o computing another divest index and is therefore jut a univariate ppreach, A, more general ts, hich Honours the urna struc ‘ure, could be eonstucted by the ANOSIM procedure (Gescrited later under multivariate techniques), comp ‘ed Between every pir of replicate ABC cures. 1 iano xr ot pra hima or serge Crk (95) Sar ste te romp ore fee Sol ewe seat cs om pce abundance atom th age ‘Soran pacha xing proba nh at ‘iat b ove meres (fo sa sets (te) ond cay at 400 aay eo ack pate Inter pansy oh nae to fare {Cameron Mins approach er eng eu of oe Imre gues diate, ba la bad ok pecs imeem mig sane strut empl ioe chine Sua fans fo fo age itelcondtion (roel pice) aria summaries (eg, CH, 0 Ts fr commonality of batons (eg chexqured fread Specie abundance dtrbution has Tonge tai” ith cbunce rite summaries of the curves fy regrsin) (Cowal see C812) ‘The distributonal/graphical techniques have been proposed specifically a8 a way of determining sires levels For the ABC method, the strongly polluted (istrbe) statis indicated if the abundance f-r- Jnmee cue falls above the biomass curve throughout its egth (eg. see the ltr pots in Fig. 1: the phen- ‘omenon is linked to the os of ageodied “linay™ Species and the rie of small portnists. Nowe thatthe ABC procedure claims to ghe an bsolae measure nthe sense that disturbance stasis ati able on the bass of simples from a single site in Practice however iti aways wise to design collection fiom (matched) impacted and contol sites o conten that the control condition exhibits the undisturbed ‘ABC pattern (biomass curve above the abundange curve, thoughout) Similarly, the species abundance distribution has features characteristic of disturbed sats (eg se the middle plts in Fig. 1.6), namely a move toa less“ shape” disteibuton by a reduction inthe first one or ‘wo abundance clases (lose of rust specs), ‘combined with the gain of some higher abundine lasses (Very numers opportunist specie). ‘The disibuionlgraphical metheds may thas have particular merits in allowing recognition of stressed States (Chapter 14), though they have the disadvant- ge of being more dificult o work with statistical for example in linking t0 environmental variables where the only viable course again seems tobe rect fon of the curve(s) for each sample 10 summary statistic (suchas 7), which ean be regressed on past- ‘colar abiotic variables, ale LA Lack Line mar). Mandan amas mann op) eto wa pyar (96-179) 1968 Species A a Taaopervenranas =O 7 Stel om oo pi eal Cg 0 aia flr yen Myre spina 2M 76 cinoma borate B09 Monacuaferapinose 1 oo ° pla idea vo ° Arasp. nd oo 0 Cortada gibb0 Geatnes 5 aca ide Oo 0 ° 1964 1968 1966 al a 7 005 oo = te 0 ea = ape 0 50” 0 ° a a 05 Geo eed ey 3 eg | hace od, im 00 2 07 ° 4 om oo ° oo ° ° 2 026 eo eo te 0 oo oo Pig 13. Lach Line and Loh Stn. po te ina ont Eh ample nm 185-1973 EXAMPLE: Loch Linnhe macrofauna Pearson (1975) describes time series of maerbenthie community srl, then over the period 1963~1973 incsve, at to sites in sea feh system on the west exat of Seoland((), Fig. 13) Pooling toa Single sample foreach of the 11 years resulted in hundance and biomass matrices of 115 rw species) ‘nd 11 columns (Samples), small part of which i shown in Table 14 Starting in 1966, palp-mill efflent was discharged 10 the sea lochs (Fae 13), "eplapd h farpaety fritraio i it 2 sal fi orator PRIMER whieh eps bdo od ona iformstn be sept shay) ara with the rate increasing in 1970 and a siniticant resistin taking place in 1972 (Pearson, 1575). The {op leftchand plot of Fig 1 shows the Shannon divers i of the macrobenthie samples over this prod, and the remaining plas the ABC curves foreach year” “There appears to be a consistent change of stucture from one in sshich the Biomass curve dominates the lndane ease inthe erly yeas fo the curves cross ing, reversing altogeter and then finally reverting {heir original fom, EXAMPLE: Garroch Head macrofauna Pearson and Blackstock (1984) describe th sampling fof a transect of 12 sites acrose the sewage-siadge

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