1982
I MARINE ECOLOGY  PROGRESS SERIES
Mar. Ecol. Prog. Ser.
I Published April 9
ABSTRACT: A strategy is presented for analysing marine biological survey data and relating the biotic
patterns to environmental data. To avoid circular argument, biotic and environmental data are kept
separate. The strategy is illustrated by a worked example using data on the distribution of 182
nematode species in 107 samples in the River Exe estuary. Nineteen stations are grouped Into 4 main
clusters using complementary classification and multidimensional scaling (MDS) ordination tech
niques. These are both based on rootroot transformed abundance data with the BrayCurtis measure of
similarity. Indicator species characterising each group are extracted using information statistics.
Inverse analyses give clusters of COoccurnngspecies which are strongly related to the station groups.
Relationships of station groups to environmental variables are revealed by superimposing data for one
variable a t a time on the MDS plot, showing that some station groups differ in sediment granulometry
and others in salinity, for example. Some of the other factors plotted show n o difference between station
groups. Similarly, physiognomic charactcrlstics of the species are superimposed on the MDS plots of
the inverse analysis of species groups, revealing differences in setal length and trophic status between
the species groups. Finally, the 4 major station groups a n d species groups are related to one another in
terms of morphological adaptation to the habitat.
O InterResearch/Printed in F. R. Germany
38 Mar Ecol. Prog. Ser. 8: 3752, 1982
suitable in pollution surveys, or when one knows in (2) Coded abundances: semiquantitative data coded
advance which physical variables are likely to be on an arbitrary scale to indicate relative abundance
dominant, but w e prefer the first approach as a general (e.g. from absent, 0, present, 1, fairly common, 2, to
procedure. In other words we analyse the biotic data dominant, 5; Field, 1970). Coding usually tends to have
first, 'letting the species tell their story' (Day et al., the effect of normalizing data and subsequent transfor
1971) and once groups of biotically similar samples mation is then unnecessary.
have been recognized we test the environmental vari (3) Frequencies: The number of occurrences of each
ables for statistical differences, an approach also species is counted, e.g. the number of grabs or hauls in
adopted by Green and Vascotto (1978) inter alia. This which the species was identified at a particular station.
strategy keeps the analyses of biotic and environmen Frequency data are often obtained by reducing a large
tal data completely separate, avoiding the influence of number of samples (hauls, cores, grabs) to a smaller
any previous assumptions about relationships between number of stations at which they were taken.
the biota and environment, and minimizing the danger (4) Densities: the commonest data, in which the
of circular argument in seeking to deduce relation number of individuals per sample is recorded.
ships. Densities often provide very skewed data, and trans
formation is normally advisable.
(5) Biomass: colonial animals and plants cannot be
METHODS meaningfully counted, whereas all organisms can be
expressed in units of weight. There is a danger of
Normal Analysis occasional, random inclusion of particularly large
organisms which may swamp the other data, therefore
For simplicity we deal first with the commonest type transformation is usually advisable.
of analysis, normal or qtype analysis, in which sam
ples (or stations) are arranged into groups which each
have a similar biotic composition. Fig. 1 summarizes Transformation
the stages of analysis.
Clifford and Stephenson (1975) discuss a number of
methods of transformation and standardization, the
Raw Data essential difference being that transformation alters
the score for each species in each sample without
The biotic data consist of a matrix in which n sam reference to the range of scores in the rest of the data. A
ples (or stations) are described by s species (or other number of transformations are commonly used, the
taxa). Data may be categorized into 5 types: commonest being the logarithmic transformation:
(1) Presenceabsence data, in which species are
recorded as being present (1) or absent (0) in each Y , = log ( X , + 1)
sample. where Xii = raw data score of the ith species in the jth
l'
Classification
\
indicator
species
Ordination
Fig. 1. Normal (qtype) analysis: diagrammatic summary of stages leading to classification and ordination of samples. Raw data
(Stage 1) are represented in a matrix of n samples by s species. It may be necessary to transform data (Stage 2). Comparison of
each sample with every other sample using a measure of similarity leads to a triangular similarity matrix (Stage 3). Classification
(Stage 4) and ordination (Stage 5) are complementary pictorial summaries of the relationships between 8 samples in this
diagram. 'Indicator species' are obtained directly from raw data. Stages are referred to by number in the text
Field e t al.: Analysing multispecies distribution patterns 39
dendrogram is arbitrary and 2 adjacent samples are not biological qtype analysis the relation is usually non
necessarily the most similar. This can be illustrated by linear, as the scatter plot for the final configuration of
visualizing a n upsidedown dendrogram as a sus the Exe estuary stations illustrates (Shepard diagram,
pended mobile with the threads holding samples Fig. 5). Nonmetric MDS allows for this by fitting a
which are free to rotate in a horizontal plane. general monotonic (increasing) regression of {dlk} on
(4) Dendrograms tend to over emphasize discon { b j k }the
, only information used from the dissimilarities
tinuities and may force a graded series into discrete being their rank order.
classes. (3) The 'goodnessoffit' of this regression is mea
In view of these disadvantages, it is advisable to sured by some criterion, usually the stress formula:
employ an additional method of presentation to show n n n n
individual relationships. If the 2 complementary
methods agree, then discontinuities can be accepted as
Stress 1 = Z: (d,,
I k>l
 4,)'/ 1 X d $
I j>k
A
real; if not, one has to temper the interpretation of where dj, = distance estimated from the regression,
results accordingly, but it may be helpful to use the corresponding to dissimilarity, If stress is large, the
distinct classes shown by the dendrogram to simplify current 'map' tallies poorly with the observed dis
description, e.g. in plotting on a map or by delineating similarities; conversely, low stress indicates that the
dendrogram classes on the corresponding ordination. sample relationships can be well represented by a
station 'map' in the specified dimensionality. Stress
may be thought of as the distortion involved in 'com
Ordination pressing' the data to a small number of dimensions.
(4) The current configuration is perturbed in a direc
Our preferred method of ordination is multidimen tion which decreases the stress (the method of steepest
sional scaling (MDS), a technique first derived for use descent is usually employed here) and stages (2) to (4)
in psychology and sociology (see Kruskal and Wish, repeated until no further reduction in stress is possible.
1978, for an introductory survey) but potentially of As with many multidimensional minimisation prob
wide application to the biological sciences. In the lems, convergence may often be to a local rather than
context of sample analysis, MDS produces an ordina global minimum of the stress function. Small perturba
tion of the n stations in a specified number of dimen tions of the present configuration may all lead to
sions. Firstly, it interprets some function of the dissimi higher stress values but an entirely different configura
larity measure between each pair of stations as a dis tion may have a lower stress minimum. It is therefore
tance in ordinary Euclidean space. It then seeks the advisable to repeat the iteration process with several
best possible reconciliation (according to some opti different starting configurations. If essentially the
mality criterion) of these n (n  1)/2 interstation 'dis same configurations (with the same lowest stress) are
tances' with the physical distances between n points achieved on a number of occasions, then the optimal
on a 2 (or higher) dimensional map. Metric MDS solution has almost certainly been found, though this
(essentially Principal Coordinates Analysis) stems can never be guaranteed. It is important to note that
from the work of Torgerson (1958); nonmetric MDS, a the final map is only determined to within an arbitrary
much more flexible tool, dates to Shepard (1962),Krus orientation and reflection, and arbitrary location and
kal (1964) and others. Kruskal (1977) reviews 2 of the scale; this explains the omission of axes in Figs. 4, 6,
more widely available computer programs for irnple and 8 to 10.
menting nonmetric MDS, namely MDSCAL and Step (2) highlights the advantages of MDS over
KYST. (The version MDSCAL 5MS has been used to techniques such as principal coordinates, reciprocal
derive the results for our example). Nonmetric MDS is averaging and correspondence analysis for obtaining a
an iterative procedure with the following steps: simple ordination. The latter are all essentially based
(1)A starting 'map' of the n stations is constructed, in on the eigenvalue method of principal components
the required number of dimensions. This could b e the (though they differ in the standardisations and trans
result of an ordination by another method, e.g. princi formations initially applied to the data) and are rela
pal coordinates analysis, but will often be simply an tively inflexible, particularly with regard to the large
arbitrary configuration chosen at random. number of zero counts generally present in a species
(2) The interpoint distances {dlk : k > j; j = l, . . . n) samples matrix, as discussed under 'Measurement of
of this configuration are then regressed on the corres Similarity'. By contrast, great flexibility is bestowed on
ponding dissimilarities {ajk).In the very simplest case, MDS by its less direct approach  first constructing a
where there is reason to believe that the dissimilarities dissimilarity matrix to suit the particular form of the
are directly proportional to distance, this could b e by data and then allowing a general monotonic transfor
simple least squares linear regression. However, for mation to distance. In fact it is surprising that such a
Field e t a1 : A n a l y s ~ n gmultlspecies d ~ s t r l b u t i o npatterns 41
precise 'map' of the relation of samples to each other tests' to pick out discriminating species). Nevertheless,
can be constructed solely from information of the type we have found it a very useful way to reexamine the
'species composition at Station A is more like that at B data in practice and recommend its use, provided it is
than at C'. not regarded as a statistical test of significance.
Further advantages of MDS are its ability to handle, Another technique that has been used to extract the
with comparative ease, missing data, replication and species that differ most between samplegroups is the
data of nonuniform reliability for which it is desirable Fratio (Stephenson et al., 1977; Shin, 1982) which can
to give unequal weights to the dissimilarities in seek be used to find the species that differ most among all
ing the 'best' map. A definite disadvantage is the the sample groups simultaneously, as compared to the
escalation of the computing problem for large numbers pairwise Information Statistic tests. The Fratio should
of stations; computer time increases proportionally to n be applied to transformed data. Similar constraints
and tends to become prohibitive for 3 figure values preclude its use a s a rigorous statistical test.
of n.
Having summarized the analysis of distribution pat Grouping of species may be of greater interest than
terns in 2 complementary diagrams, classification and samplegroups in which case inverse or ranalysis is
ordination, one has lost track of the species differences appropriate a s a complen~entor substitute for analysis
which cause the patterns. This information can b e of samples.
regained by returning to the raw data, or preferably to If presenceabsence data are to b e analysed either
frequencies derived from the raw data. the BrayCurtis measure or that of McConnaughey
It may be useful to know which species are charac (1964) have proved satisfactory (Field, 1970), but to
teristic of one group of samples but absent from avoid spurious groups of rare species the number of
another. Information statistic (I) tests (Field, 1969; species should b e reduced. If semiquantitative or
Velimirov et al., 1977) provide a means of assessing quantitative data are used, the BrayCurtis measure is
which species differ most between one group of sam appropriate only if the data are first standardized (W.
ples and another. Comparing Cluster 1 with Cluster 2 Stephenson, pers. comm.).
for any one taxon (species), i: A disadvantage of inverse analysis alone is that later
analysis of the species groups in relation to the
environmental data is more complex and simple tests
where I,, = total information content of both clusters for significant differences between groups are not
combined, I , = N, log N,  A,, log A,, (NrAti) log (N,  appropriate. This can b e partially remedied i n a pre
A,); N,= number of samples in both clusters together senceabsence analysis by contradicting our overall
('potential presences'); A, = number of samples in strategy and categorizing the environmental data, the
which Species i is actually present; (N, A,) = number classes of which can b e treated as 'species' which are
of samples from which Species i is absent. Similarly, 'present' or 'absent' from samples a n d grouped into
the information content I,, and I,, are obtained for clusters along with the biotic species (Field, 1971).
Clusters 1 and 2 respectively. Thus any clusters may be Steps 25 are followed as with normal analysis, but
compared, pairwise, to s e e which species discriminate additional stages (7) and (8) are also required (Fig. 2).
best between them.
Under the hypothesis that the probability of observ
ing Species i is the same for samples in both Clusters, Data Reduction
2 A 1 , has a n approximate chisquare distribution, and
the scores of each species might be looked up in chi In any survey, some species occur too seldom to form
square tables (1 d.f.) to s e e whether they differ signifi a n analysable pattern. These a d d to computer time but
cantly at the 5% or 1 % probability levels. However, they do not affect normal analysis of samples (Day e t
these are best regarded a s arbitrary cutoff levels, a al., 1971). However, these problems become more seri
convenient rule of thumb for selecting indicator ous when species a r e compared (inverse analysis) for 2
species but lacking statistical rigor. This is because reasons. Firstly, computer time increases with the
several assumptions implicit in chisquare testing are square of the number of individuals (now species)
not met (e.g. large sample conditions such as expected being compared; secondly, random COoccurrenceof 2
values exceeding 5, repeated significance tests, and rare species which only appear once in the analysis,
the fact that classification categories have already can result in their being grouped together as having
partly been determined by the same data used in 'I identical distributions.
Mar. Ecol. Prog. Ser. 8 . 3752, 1982
U
Classification
Similarity
I *
Ordination
Matrix
Fig. 2. Inverse (rtype) analysis: diagrammatic summary of stages leading to classification and ordination of species. Additional
steps (7 and 8), reduction of the number of species and standardisation of abundances respectively, are usually required before
comparing each of the S' species with every other species. The classification suggests that there are 2 main clusters of 4 species
each; the ordination indicates which individual species are close or distant in distribution
Clifford and Stephenson (1975) discuss several pos sample; C X,, = summed abundance of the ith species
,=l
sible ways of eliminating rare species, and Stephenson over all samples; Y,, = corresponding standardized
and Cook (1980) describe a method of eliminating score. This type of standardisation is also referred to as
species prior to analysis. In many analyses, as in the 'relativized data' (Whittaker and Gauch, 1973; Camp
example below, some stations will tend to have low bell, 1978).
species diversity but high numerical abundance, and
others a high diversity and low abundance. Thus, to Relationship to Environmental Data
select say the 50 overall most abundant species, biases
the selection towards the species at the low diversity1 Having obtained groups of samples by analysis of
high abundance stations whereas many species which the biotic data, most ecologists are interested in seek
may be absolutely characteristic of the high diversity1 ing the environmental factors that are likely to be
low abundance stations will be disregarded. To over responsible for the patterns found. One approach is to
come this problem, we have selected on the basis of do this in a separate statistical analysis of the environ
species having above an arbitrary percentage domi mental data.
nance at any one station. A variety of techniques is available for this, ranging
from simple ttests or their nonparametric counterpart,
MannWhitney Utests (Siegel, 1956) to analysis of
Standardisation variance and multiple discriminant analysis (Polgar,
1975; Green and Vascotto, 1978).
NoyMeir (1970, 1973) has discussed fully various In the simplest case, 2 samplegroups are compared.
types of standardization used in plant ecology, and a All observations of an environmental variable (e.g.
summary is given by Clifford and Stephenson (1975). temperature) pertaining to one sample group are
For normal analysis, standardization is not required tested against the corresponding observations of the
but it is necessary prior to inverse (species) compari other sample group. This is done using each suggested
sons when quantitative data are used. This is because variable in turn and the ones that differ significantly
perfectly correlated species which always occur are noted as being possible factors responsible for the
together (e.g. host and parasite) but in different abund biotic groups (Field, 1971).This approach has the usual
ances might be separated from one another because drawbacks inherent in repeated significance tests and,
their scores are not the same. The recommended stan because only one pair of samplegroups can be com
dardisation for species comparison is: pared at a time, the overall effect of environmental
parameters on the observed patterns is not clear. With
Y, = 100 X,] / l 1
=1
X,] ordination techniques it is often found that the samples
or stations assume configurations which are orientated
where X,, = abundance of the ith species in the jth along particular dominating environmental gradients.
Field et al.: Analysing multispecies distribution patterns 43
In the worked example below we have therefore given in Warwick (1971). In general the muddy sta
ranked several environmental parameters and plotted tions had a high population density but low species
them on the station configurations of the ordination to diversity and the sandy stations the reverse. The
give a visual impression of possible correlations. MHWST sandy stations at Lympstone and Shelly Bank
had a low diversity/low density fauna, attributed by
Warwick (1971) to seepage of low salinity coastal sub
A Worked Example soil water.
% SIMILARITY
Fig. 3. Dendrogram showing classification of 19 stations in Exe estuary based on mean bimonthly abundances of nematodes over
1 y. Abundances were rootroot transformed before comparing stations using the BrayCurtis measure, and the dendrogram
formed by groupaverage sorting. Four main station groups ( 1 4 )are distinguished at an arbitrary similarity level of 15 % (Xaxis)
Further discussion will only concern the station Table 2 . Frequencies of occurrence of indicator species,
means data, since no seasonal differences are evident. ranked according to information statistics, which distinguish
stations of Group 1 from those of Groups (2 + 3). Rigorous
statistical criteria are not met; species tabulated above the
arbitrary horizontal line have 2 A I > 6.63, those below the
Indicator Species line have 2 D I > 3.84. Number of occurrences in Group 1 are
given, with those in Groups (2 + 3) in parentheses; maximum
Tables 2 and 3 list the species which are characteris values are N 1 = 7 and N (2 + 3) = 4, respectively
tic of Group 1 and distinguish i t from Groups (2 3) +
Species Group 1 (Groups 2 + 3)
Mesothenstus setosus
Desmolaimus zeelandicus
Leplolaimus papilliger
Table 3. Frequencies of indicator species, ranked according to Table 5. Frequences of occurrence of indicator species,
information statistics, which distinguish stations of Group 1 ranked according to information statistics, which distinguish
from those in Group 4. Numbers of occurrences in Group 1 are Group 4 from Groups (2 + 3) There are no 'perfect indicators'
given with those in Group 4 in brackets; maximum values are present in all 8 stations of Group 4 and none of Groups (2 + 3)
N 1 = 7 and (N 4 = 8), respectively
Species N4 = 8 ( N [ 2 + 3 ] )= 4 )
Species Group 1 (Group 4)
' Sigmophora litoralis 7 (0)
Mesotheristus setosus 7 (1) ' Ditlevsenella danica 6 (0)
Desmolairnus zeelandicus 7 (1) Theristus sp. G 6 (0)
Sabatieria pulchra 7 (2)
Cylindrotheristus oxycercus 7 (2) ' Alaimella truncata 5 (0)
' Leptolaimus papilliger 5 (0) Epacanthion gorgonocephalum 4 (0)
Anoplostoma viviparurn 7 (3) ' Chromaspirina parapontica 4 (0)
Axonolaimus paraspinosus 4 (0) Oncholaimellus calvadoscus 4 (0)
Thalassironus sp 4 (0)
Viscosia viscosa 5 (1)
Tripyloides gracilis 3 (1) This species comprises less than 4 % of numbers at any
Sphaerolairnus hirsutus 3 (0) one station and is excluded from the inverse specles
Spilophorella para doxa 3 (0) analysis (Table 6)
Odontophora setosa 3 (0)
Oxystornatina elongata 3 (0)
Theristus procerus 3 (0)
This species comprises less than 4 % of numbers at any Inverse Analysis
one station and is excluded from the inverse species
analysis (Table 6) For the species analysis we have reduced the total
numbers from 182 to 55 by using only those species
which have > 4 % dominance at any one station.
of the species listed in Tables 2 to 5 may b e useful Species abundances have been standardised for each
indicators, but are not sufficiently dominant to be station as a percentage of the total abundance at all
included in the inverse analysis which follows. It stations (i.e. if a species is found at only 1 station, its
should be noted that the technique is most effective abundance there is 100%). Fig. 7 shows the dendro
when the 2 groups being compared are both large, gram for the inverse analysis. In order to define the
hence w e combined Groups 2 a n d 3 in the compari same number of species groups as station groups w e
sons. have drawn horizontal lines at 6 % similarity and at
8 % similarity giving 5 groups (if station groups 1A and
Table 4. Frequencies of indicator species, ranked according to 1 B are considered separately). For convenience the
information statistics, which distinguish Group 4 from species clusters on this dendrogram have been desig
Group 1. There are no 'perfect indicators' present in all 8
nated by the same notation as the station clusters, since
stations of Group 4 and absent from all 7 stations of Group 1
they clearly correlate with them (see below). A list of
the species in each cluster is given in Table 6. There is
Species N 4 = 8 (N1=7)
good general agreement between the species groups
listed in Table 6 and the indicator species extracted by
' Sigmophora litoralis 7 (0)
' Ditlevsenella danica 6 (0) the information statistic technique. Five out of 6
Ther~stussp. G 6 (0) species characterising Group 1 in Table 2 are in
' Alaimella truncata 5 (0) Group 1 of Table 6, the 6th being too rare for inclusion
in the inverse analysis. Similarly, all but 1 of the
Epacanthion gorgonocephalum 4 (0)
included species characterising Group 1 in Table 3 are
' Chromaspirina parapontica 4 (0)
0nchola1mellus calvadoscus 4 (0) also clustered into Group 1 by the inverse analysis;
' Thalassironus sp 4 (0) Tripyloides gracilis is included in Group 2 (Table 6).
' Leptonemella sp. 3 (0) However, only Epacanthion gorgonocephalum and
' Linhomoeus sp. 3 (0) Oncholaimellus calvadoscus are included in Group 4
' Theristus albigensis 3 (0)
' Trichenoplus sp.
by the inverse analysis (Table 6) and also distinguish
3 (0)
' Linhomoeus sp. B 3 (0) Group 4 from Groups 1 , 2 and 3 (Tables 4 , 5). Thus it
appears that a number of rarer species occur only in
This species comprises less than 4 % of numbers at any
one station and is excluded from the inverse species Group 4, and these are too rare to appear in the inverse
analysis (Table 6) analysis.
Fig. 8a is the species analysis with multidimensional
F ~ e l det al.: Analysing multispecies distribution patterns 4
Fig. 7. Dendrogram of inverse analysis comparing 55 nematode species occurring in more than 4 % dominance at any of the 19
River Exe stations. Species abundances standardised and compared using the BrayCurtis measure with groupaverage sorting.
Species numbers listed and species named by cluster in Table 6
Table 6. Species groups distinguished by inverse (rtype) analysis. The groups are based on the dendrogram in Fig. 8 ; species
numbers refer to numbers used in Fig. 7 and 8 Nomenclature follows that of Gerlach and Riemann (1973,1974)
Group 1A Group 3
1 Mesotheristus setosus; 3 Sabatieriapulchra; 5 Hypodon 15 Tripyla so.; 16 Rhabditid; 17 Dorylaimid; 18 Eury
tolaimus geophilus; 2 Anoplostoma viviparum; 6 Desmo stomina terricola; 14 Bathylaimus stenolaimus; 35 Paracy
laimus zeelandicus; 4 Axonolaimus spinosus; 8 Adon atholaimus intermedius
cholaimus thalassophygas; 1 1 Theristus flevensis
Group 4
Group 1B 30 Enoplus brevis; 37 Mesacanthion africanthiforme; 38
12 Axonolaimus paraspinosus; 24 Paracanthonchus punc Enoploides brunettii; 39 Enoplolaimus litoralis; 41 Trisson
tatus; 9 Viscosia viscosa; 10 Ptycholaimellus ponticus; 7 chulus benepapillosus; 42 Araeolaimus elegans; 48
Cylindotheristus oxycercus; 25 Sphaerolaimus hirsutus; 22 Epacanthion gorgonocephalum; 49 Enoplolaimus
Odontophora setosa; 33 Atrochromadora microlaima; 34 denticulatus; 54 Axonolaimus hexapilus; 31 Chromadora
Terschellingia communis nudicapitata; 44 Mesacanthion hirsutum; 40 Enoplolaimus
propinquus; 43 Dichromadora hyalocheile; 45 Praeacan
Group 2 thonchus opheliae; 46 Viscosia cobbi; 47 Sigmophora
20 Paracanthonchus tyrrhenicus; 21 Tripiloides gracilis; 13 rufum; 53 Bathylaimus paralongisetosus; 50 Pomonema
Oncholaimus brachycercus; 19 Ascolaimus elongafus; 22 reducta; 51 Oncholaimellus calvadoscus; 52 Axonolaimus
Adoncholaimus fuscus; 23 Theristus acer; 26 Microlaimus orcombensis; 55 Chromaspirina inglisi
honestus; 29 Microlaim us robustidens; 32 Trefusia lon
gicaudata; 28 Theristus normandicus; 36 Bathylaimus
assimilis
scaling; the species groups are delineated from the cates a high degree of agreement between the species
dendrogram. Fig. 8b shows the same configuration and station clusters. Although the classification artifi
with the species numbers replaced by the station cially forces a species to occur in only 1 group, it will
group(s) in which they are found to represent more be seen from Fig. 8b that, in this case, rather few
than 4 % of specimens in any one station. This indi species occur in significant numbers in more than 1
48 Mar. Ecol. Prog. Ser 8 . 3752, 1982

KEY
LINEAR SCALE FOR LINEAR SCALE FOR
AVERAGE SAUNITY M A N DEPTH OF
~ A S B O F ~ H S L AYER. E.G.
2
SLAWTER). L G
EXCEPT
I >'OCM"
Fig. 9. Relation of station
groups to environmental fac
tors. MDS plot of 19 River Exe
W 1151.
LINEAR SCALE FOR
stations based on nematode
MEDIAN D w m R MLWST fauna with clusters delineated
OF PARTICLES. EG. as in Fig. 4. (a) At each station
0 MLWNT
0 0.2 MM. circles proportional in diame
0 MTL ter to mean salinity of intersti
0 0.5 mm.
tial water are superimposed.
0 MHWNT
01.0 MM. (b) Relation of station groups
0 MHWST to particle size, with circles
proportional in diameter to av
erage median sediment parti
cle size at each station. (c) Re
lation of station groups to sedi
ment organic content: circle
KEY diameters represent mean %
S Q U A R E ROOT organic content at each station
SCALE FOR MEAN
%ORGANIC
on a square root scale. (d) Re
CONTENT. E.G lation of station groups to
depth of H,S layer in sedi
ment: arrow lengths propor
tional to mean depth of H,S
layer at each station. (e) Rela
tion of station groups to tidal
level: columns indicate height
of each station above chart
datum

KEY 
KtV
SCTAL LENGTH o,M>
RECER
l 510
bodied species, but there is a suggestion that Group 4 enable them to maintain their position in this highly
has proportionally fewer small (or more large) species. dynamic environment. Stations 1219.
(4) Cuticle pattern (Fig. 10d): No marked difference in Although these conclusions broadly agree with the
the distribution of cuticle patterns between groups. subjective assessment made earlier (Warwick, 1971),
The statistical significance of the differences noted they differ somewhat in detail. Warwick did not note
above can be examined by a chisquare test. For a 2 the separation of Group 1 stations into 2 coherent sub
way categorisation into 'feeding type' and 'species groups on the basis of salinity, and included Station 8
cluster', a test of the null hypothesis that there is no from Group 1B with Group 2 stations because of its
association between the classifications is just rejected slightly coarser sediment. The present study shows
at the 5 % level, whereas independence of 'cuticle quite clearly that, faunistically, Station 8 belongs with
pattern' and 'species cluster' cannot be rejected. The Group 1B. Further, Warwick divided the Group 4 sta
continuous variables can be categorised and a chi tions into 3 groups: Stations 12 and 13  coarse sands
square test applied similarly; 'setal length' is seen to which dry out at low tide, Stations 14 to 18 coarse sands
b e very strongly associated with species group ( x 2 r= with a more or less permanent water table, and Station
29.6 on 9 degrees of freedom), but the suggestion that 19 fine stable sand retaining a permanent water table.
'body length' differs between the clusters is not strong The present study suggests that there was no basis for
enough to attain statistical significance. this separation and that the water table or the precise
As an alternative to categorisation, the observation grade of sand have little effect on the clustering of
that setal length tends to increase along in axis from stations within Group 4. Warwick's observations on the
bottom right to top left of the MDS plot can formally be distribution of physiognomic characters, bearing in
verified by multiple linear regression. For the regres mind some regrouping of stations, are largely borne
sion of log setal length of the two MDS coordinates, out by the present study.
the slope of the fitted plane is in the anticipated direc
tion and is very significantly nonzero (F = 12.5 on
2.52 degrees of freedom). For body length regressed in General Conclusion
the same way, the slope does not achieve significance,
in agreement with the earlier x2test. Application of our strategy has given as a first result
a dendrogram which divides the stations into 4 groups,
one of which is subdivided. The simplicity of the clas
Main Conclusions from Worked Example sification is useful for presentation but may force the
data into artificially distinct classes when continua
These analyses show that, in the Exe estuary, there exist, hence a complementary method of analysis is
are 4 major groups of stations representing, in physico advisable. Multidimensional scaling (MDS) con
chemical terms, 4 major habitats each characterised by firmed the existence of clear groups which were
a set of nematode species which have morphological emphasized by delineating the dendrogram groups on
adaptations to suit the habitat. the MDS plot. Inverse MDS analysis (after excluding
(1) Muddy stations with a high organic content and a the rarer species to reduce computer storage and time)
blackened anoxic layer a few cm below the surface, produced clusters of species which correspond well
characterised typically by small depositfeeding with the station groups. Information tests were used to
nematodes with short setae. Salinity either low (Group find 'indicator' species characteristic of one group of
l A , Stations 14) or moderate (Group lB, Stations 79). stations and absent from another group.
(2) Muddysand stations at MHWNT with moderate The second stage of analysis, according to our
salinity and organic content with blackened H2S layer strategy, is to relate to environmental data the groups
at variable depth. Physiognomic characters of the formed by analysing the biotic data. This was done by
nematodes intermediate between mud (Group 1) and superimposing a scaled symbol, representing one
sand (Group 4) stations. Stations 6 and 11. environmental variable, onto the MDS plot of the sta
(3) Welldrained sands at MHWST with moderate tions obtained previously. This demonstrated clearly
organic content, very low interstitial salinity and no whether a factor, such as salinity, differs markedly
H2S. Rather few species so that the distribution of between the station groups. We could have used statis
physiognomic characters cannot be considered, but 3 tical tests (e.g. ANOVA or KruskalWallis tests) to test
of the 6 species in this group belong to freshwater the significance of differences in each environmental
genera. Stations 5 and 10. variable among the station groups; this was unneces
(4) Clean sands, high salinity, no H2S, low organic sary in our worked example because of the very
content. A high proportion of large predatory or marked differences.
omnivorous nematode species with long setae which Finally, the same graphical approach of superimpos
Field e t al.: Analysing multispecies distribution patterns 51
ing s y m b o l s o n t h e MDS p l o t s can b e u s e d t o s h o w list of aquatic nematodes (Part 2). Veroff. Inst. Meeres
r e l a t i o n s h i p s b e t w e e n a t t r i b u t e s of t h e s p e c i e s ( h e r e forsch. Bremerh. 2 (Suppl. 4): 405734
feeding type a n d morphometry) o n the o n e hand, a n d Green, R. H., Vascotto, G. L (1978).A method for the analysis
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This paper was submitted to the editor; it was accepted for printing on January 3, 1982