Journal of Animal

Ecology 0888\
57\ 658671

The taxonomic distinctness of coastal bottom!dwelling

_sh communities of the North!east Atlantic
STUART I[ ROGERS\ K[ ROBERT CLARKE$ and
JOHN D[ REYNOLDS%
CEFAS\ Lowestoft Laboratory\ Pake_eld Road\ Lowestoft\ NR22 OHT\ UK^ $CCMS\ Plymouth Marine
Laboratory\ Prospect Place\ West Hoe\ Plymouth\ PL0 2DH\ UK^ and %School of Biological Sciences\
University of East Anglia\ Norwich\ NR3 6TJ\ UK

Summary
0[ New techniques for identifying the average taxonomic range of species assemblages
were applied to an extensive dataset of bottom!dwelling _sh in the coastal waters of
NW Europe[ These taxonomic distinctness indices provided much greater resolution
than traditional diversity indices as they incorporated information on taxonomic
relationships into an index which measures species dominance[ Unlike standard mea!
sures of species richness and diversity\ the mean value of these statistics is independent
of sampling e}ort\ and this allows objective comparisons to be made between samples
from studies where sampling e}ort is not standardized[
1[ A reduction in the average taxonomic range between the fauna of western waters
of the UK and that of the southern North Sea was consistent with the general decline
in species richness observed between these regions\ and suggests that these two factors
may be spatially positively correlated[ Indices calculated for individual samples of
_sh on a local scale\ however\ did not all _t this trend[
2[ Much of the variability in taxonomic diversity within the coastal waters of NW
Europe was caused by the variable geographical distribution of the elasmobranchs[
Of all the families which comprise the _sh communities\ this group has life!history
characteristics which make it most susceptible to impact by commercial trawl _sheries[
3[ The use of taxonomic distinctness measures provided additional insights\ of rel!
evance to biodiversity assessment\ suggesting that they might usefully be applied to
other aquatic and terrestrial fauna[
Key!words] taxonomic diversity\ ground_sh\ community structure\ _shing impact\
NW Europe[
Journal of Animal Ecology "0888# 57\ 658671

Introduction
Explaining the large!scale patterns in species dis!
tribution and abundance of both marine and ter!
restrial environments requires information on habitat
extent and complexity "Magurran 0877^ Rosenzweig
0884#\ the rates and e}ects of disturbance and levels
of productivity "Connell 0867^ Sousa 0868^ Petraitis\
Latham + Niesenbaum 0878^ Baltz 0880#\ and also the
impact of environmental gradients on species richness
"Pianka 0855^ Ekman 0856^ Nelson 0883#[ Rarely are
there su.cient environmental data to be able to fully

0888 British
Ecological Society

Correspondence] Dr S[ I[ Rogers\ CEFAS\ Lowestoft Lab!

oratory\ Pake_eld Road\ Lowestoft\ NR22 OHT\ UK[ Fax]
90491 402754[ E!mail] s[i[rogerscefas[co[uk[

explain patterns on a smaller provincial or regional

scale\ but it has become increasingly necessary to at
least understand the relative role of natural and
anthropogenic perturbation[
The response of benthic marine assemblages to dis!
turbance is thought to be more easily detected at
higher taxonomic levels\ because of the confounding
in~uence of abiotic factors such as water depth and
temperature which will a}ect small!scale diversity at
the species level "Warwick + Clarke 0882#[ This hier!
archical response to stress\ _rst proposed by Pearson
+ Rosenberg "0867#\ exploits the fact that various
phyla di}er in their sensitivity to disturbance[ Benthic
environments which have been perturbed are gen!
erally kept in an early successional stage with low
species diversity\ and often consist of species which
658

669
Taxonomic
distinctness of _sh
communities

0888 British
Ecological Society
Journal of Animal
Ecology\ 57\ 658671

are closely related\ while unperturbed benthic com!

munities in a late successional stage often consist of a
wider range of taxonomically distinct species "War!
wick + Clarke 0884#[ It is not clear\ however\ whether
all communities which have a small number of species
should\ by de_nition\ have a more limited taxonomic
range than those with many species[ Assuming that
taxonomic diversity translates into ecological diver!
sity\ then the taxonomic range of an assemblage may
be crucial in maintaining ecosystem stability during
natural or anthropogenic perturbations "Hughes
0883^ Tilman 0885#[
Obscuring these patterns are the underlying bio!
geographic processes\ such as rates of speciation\
immigration and extinction\ which determine total
species richness of a province "Rosenzweig 0884#[ The
resulting species diversity gradients are largely deter!
mined by the in~uence of environmental factors on
rates of diversity generation and extinction[ For mar!
ine organisms\ the species richness between the tropics
and polar regions generally\ although not always\
shows consistently decreasing trends "Findley + Find!
ley 0874^ Pauly 0883# but see also Gee + Warwick
"0885#[
These observations in~uence the way we interpret
species diversity trends in marine environments\ yet
the conventional indices of community diversity\
which use only the relative abundance of species\ do
not describe the degree of taxonomic relatedness of
those species[ Assemblages with the same species rich!
ness may either comprise species which are closely
related to one another taxonomically\ or they may
be more distantly related "Ludwig + Reynolds 0877^
Warwick + Clarke 0884#[ To quantify such changes
in taxonomic relatedness\ Warwick + Clarke "0884#
and Clarke + Warwick "0887b# have de_ned three
biodiversity indices\ which quantify the taxonomic
diversity and taxonomic distinctness of a faunal
assemblage using the path lengths between _sh gro!
uped by their taxonomic relationships[ They show
that taxonomic diversity "D# is a natural extension
of Simpson diversity\ incorporating information on
taxonomic relationships within a sample into an index
measuring species dominance[ Taxonomic dis!
tinctness "D# is a form of taxonomic diversity that
limits the in~uence of patterns in species dominance
by dividing D by a form of Simpson diversity\ thus
constructing a measure which more nearly re~ects
pure taxonomic relatedness[ A third index\ taxonomic
distinctness "D#\ considers only the special case where
abundance information is not available or is ignored
"i[e[ presence:absence data#[ Clarke + Warwick
"0887b# show that these statistics are largely inde!
pendent of sampling e}ort\ which makes their use
attractive in spatially extensive or long time!series
studies where total sampling e}ort in di}erent areas
or at di}erent times is rarely standardized[ Standard
diversity indices such as species richness\ Shannons|
index and evenness measures can be highly in~uenced

by di}ering sample sizes\ making meaningful com!

parisons impossible "Rogers et al[ 0888#[ In addition\
the presence of a statistical testing framework for the
D index enables a comparison to be made between
an observed taxonomic distinctness measure and its
expected range of variation[
These measures of taxonomic distinctness have
been applied to macrobenthos samples from the Eko!
_sk oil!_eld in the North Sea "Warwick + Clarke
0884#[ This study identi_ed a high degree of sensitivity
to subtle environmental impacts at this site\ although
these patterns have not been supported by data from
other North Sea oil _elds "Somer_eld\ Olsgard + Carr
0886#[ Taxonomic distinctness measures have also
been applied to literature data on marine benthic
nematodes from various intertidal:subtidal and coas!
tal:estuarine sites in the UK\ and also coastal habitats
in Chile "Warwick + Clarke 0887#[ These analyses
demonstrate a decrease in taxonomic distinctness D
at polluted sites\ such as the UK|s Clyde Estuary and
Liverpool Bay\ compared with taxonomic distinctness
at {clean| sites in the Exe Estuary\ the Scilly Isles and
the Northumberland coast[
The majority of published studies in which these
indices have been applied have used data from soft!
sediment macro!and meiobenthic faunas\ yet some
of the most extensive marine datasets describe the
temporal and spatial abundance of _sh[ In the only
known application of these indices to vertebrate popu!
lations\ Hall + Greenstreet "0887# show that\ for a
long time!series dataset for bottom!dwelling _sh from
the northern North Sea\ trends in taxonomic dis!
tinctness and diversity were identical to those shown
by conventional indices[ Their observed positive cor!
relation between D and Hills N0 and N1 diversity
indices suggests that assemblages that are more
diverse "species!rich# always contain species that have
a wider average taxonomic range than assemblages
which are less diverse\ but this has yet to be tested at
a range of spatial scales[
In contrast to these large!scale patterns in species
diversity\ little is known about the processes which
structure the diversity of smaller regional environ!
ments[ In this paper we re!examine an extensive data!
set describing the relative abundance of bottom!dwell!
ing _sh in the coastal waters of NW Europe "Rogers
et al[ 0887\ 0888# to test whether these new taxonomic
diversity indices can mirror changes in species diver!
sity at di}erent scales\ ranging from the entire study
area to individual sample sites[ Using a comprehensive
taxonomy of the bottom!dwelling _sh found in the
North!east Atlantic\ we _rst calculate indices of taxo!
nomic diversity using _sh relative abundance\ and also
using simple presence and absence data[ We then test
whether these taxonomic diversity indices correspond
to known patterns of species diversity\ at intermediate
and at small spatial scales[ We also describe how local
factors which may structure the community\ such as
the e}ect of man made disturbance on sensitive taxa\

660
S[I[ Rogers\
K[R[ Clarke +
J[D[ Reynolds

can in~uence the diversity of the entire demersal

fauna[

Materials and methods

SURVEYS

A series of international beam trawl surveys of north!

east Atlantic coastal waters have sampled the abun!
dance of coastal "09019 m depth# bottom!dwelling
species since the late 0879s\ using sampling techniques
described in detail by Rogers et al[ "0887#[ Di}erent
beam trawls were used during the surveys\ depending
on the ability of the di}erent vessels to deploy them\
and the varying nature of the seabed in di}erent parts
of the region[ All samples were collected during the
third quarter of the year\ at which time most _sh were
dispersed and not undergoing migrations to or from
spawning grounds[ In general\ _shing stations sam!
pled were at _xed positions\ except in Dutch and Ger!
man surveys where the station positions were strati_ed
by International Council for the Exploration of the
Sea "ICES# rectangle "0 degree of longitude 9=4
degree latitude# and selected annually on a pseudo!
random basis[ In the North Sea\ all stations were
grouped by quarter ICES rectangle\ and the centre of
this unit was used as the nominal _xed station
position[ The duration of each tow was either 04 or
29 min[ At each station the _n!_sh catch was identi_ed
to species where possible\ measured\ and their num!
bers recorded "Rogers et al[ 0887#[ Mean catch rates
of each species or taxa "raised to the number per 7 m
beam trawl per hour tow# at each _xed station position
between 5 m to 24 m depth\ and for the period 0889
85\ were calculated[ Only those _sh species which were
classi_ed as bottom!dwelling species\ and were there!
fore well sampled by beam trawls\ were used in analy!
ses of community structure "Rogers et al[ 0887#[ The
ICES Divisions surveyed were subdivided into nine
coastal sectors "Fig[ 0#[
TAXONOMIC DIVERSITY INDICES

A taxonomy of bottom!dwelling _sh species was used

to determine their relatedness "Fig[ 1#[ We compiled a
composite taxonomy based primarily on Nelson
"0883#\ with additional information for Chon!
drichthyes based on McEachran + Miyake "0889# and
McEachran\ Dunn + Miyake "0885#[ In addition to
the _ve taxonomic levels shown in Fig[ 1\ we included
subgenera\ tribes\ subfamilies\ superfamilies\ sub!
orders\ series\ superorders\ and subdivisions where
possible[ Three biodiversity indices de_ned by Clarke
+ Warwick "0887b# were then calculated from the
bottom!dwelling _sh abundances]
0888 British
Ecological Society
Journal of Animal
Ecology\ 57\ 658671

D  SSij vijxixj:n"n 0#:1

eqn 0

D  SSij vijxixj:SSdij xixj

eqn 1

D  SSij vij:s"s 0#:1

eqn 2

where xi denotes the abundance of the ith of s species

observed\ n "Sixi# is the total number of individuals
in the sample and vij is the weight given to the path
length linking species i and j in the taxonomy[ Taxo!
nomic diversity "D# can be thought of as the average
path length between any two randomly chosen indi!
viduals from the sample "including individuals of the
same species#\ whereas taxonomic distinctness "D# is
the average path length between two randomly chosen
individuals\ conditional on them being from di}erent
species[ When only presence:absence data are used
"i[e[ xi 0 0 for all species present# both equations
reduce to an even simpler form of taxonomic dis!
tinctness\ D\ which can be thought of as the average
path length between any two randomly chosen species
present in the sample[ All three indices were calculated
for the ground_sh data\ including an additional modi!
_ed form of eqn 1 in which the abundances were
square root transformed\ to downweight the con!
tribution to the index of the numerically dominant
species[
Following Warwick + Clarke "0884#\ the simplest
form of path length weighting was adopted for the
02 taxonomic levels "Table 0#\ namely] v  9 for two
individuals of the same species^ v  0 for two indi!
viduals in the same subgenus but of di}erent species^
v  1 for species in the same genus but di}erent
subgenera^ v  2 for species in the same tribe but
di}erent genera\ etc[ Where _ner subdivisions do not
exist within some groups of taxa\ the path length is
de_ned as that for the _rst existing link[ For example\
if a particular genus does not possess subgenera then
all links between its constituent species carry a weight
of v  1[
There is a degree of arbitrariness about a constant
path length of one unit between each taxonomic level
in the hierarchy^ for example\ if a wholly unused level
is inserted\ the relative distinctness values for the set of
samples will change[ Where there is a comprehensive
regional list for a group of species\ it is possible to
re_ne the weighting to re~ect the reduced number of
representatives at each taxonomic level[ An alter!
native weighting assessed here ""vo#\ Table 0# sets the
path length from one group to the next coarsest classi!
_cation as proportional to the percentage by which
taxon richness decreases[ The path lengths are then
summed and the _nal path lengths scaled so that the
total weight linking two species that belong to di}er!
ent subdivisions "the top category of the hierarchy# is
set equal to 099[ It can be seen from Table 0 that
this alternative weighting achieves the desired balance[
For example\ from genus "s2  61# to tribe "s3  56#
now adds only 2=5 to the path length whereas that
from suborder "s7  22# to order "s8  03# adds 09=4\
and addition or deletion of an unused category will
now have no e}ect at all[ The relative taxonomic dis!
tinctness indices are computed and compared for both
v and vo forms of weighting[
When the data are reduced simply to pres!

661
Taxonomic
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communities

Fig[ 0[ The shelf seas of continental NW Europe\ showing the 49 m depth contour and the nine coastal sectors used in these
analyses] "0#  Bristol Channel^ "1#  Western Irish Sea^ "2#  Eastern Irish Sea^ "3#  Western Channel^ "4#  North!eastern
Channel^ "5#  South!eastern Channel^ "6#  South!western North Sea^ "7#  South!eastern North Sea^ "8#  East Central
North Sea[

0888 British
Ecological Society
Journal of Animal
Ecology\ 57\ 658671

ence:absence\ not only can distinctness D still be

calculated\ and compared across samples of di}erent
size\ but a signi_cance test can also be carried out[
This tests for the departure of Dm\ the distinctness
measure for any sample of m species\ from the overall
value D for a {global| species list for that region[ The
test is based on the theoretical mean and variance of
Dm\ values obtained by random sampling of m species
"without replacement# from the total list of s species
"Clarke and Warwick 0887b#[ Although the theor!
etical mean remains constant\ the variance naturally
increases as m decreases\ and so the approximate 84)
con_dence intervals take the form of a {funnel|[ The
values of D for any particular set of samples can then
be related to this con_dence {funnel|\ to gauge the
extent to which their taxonomic distinctness falls sig!
ni_cantly below "or above# that expected[ Assuming a
null hypothesis that each sample is a random selection
from the total species list\ all values of D should fall
within the con_dence funnel[

From its construction\ the taxonomic distinctness

index D should be independent of the Simpson spec!
ies diversity index\ i[e[ they must be quantifying
di}erent aspects of diversity[ It is thus interesting to
show how these two community attributes are cor!
related across the available\ spatially dispersed\ sets
of samples[ Simple scatter plots are used to examine
this[

Results
The taxonomic diversity of the untransformed _sh
abundance matrix showed similar patterns for the
three diversity indices D\ D and D "Fig[ 2#[ Although
most coastal sectors showed similar values\ the East!
ern Central and South!eastern North Sea\ and the
Western Irish Sea had lower mean values of D "taxo!
nomic diversity# than others[ Mean values of D "taxo!
nomic distinctness# maintained the same general pat!
tern "Fig[ 2b#[ The use of square root transformed

662
S[I[ Rogers\
K[R[ Clarke +
J[D[ Reynolds

Fig[ 1[ The taxonomy compiled for this analysis\ simpli_ed to show _ve from a total of 02 levels of classi_cation[
0888 British
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Journal of Animal
Ecology\ 57\ 658671

663
Taxonomic
distinctness of _sh
communities

0888 British
Ecological Society
Journal of Animal
Ecology\ 57\ 658671

Table 0[ The 02 levels of classi_cation "k# used in the bottom!

dwelling _sh taxonomy\ with the number of each type "sk#\
the {standard| weighting scheme "vk#\ and an alternative
version "vko# utilising an accumulation of the proportional
decrease in taxon richness values "sk#[ More precisely\ for a
speci_c k\ vk is the weighted path length between species
belonging to di}ering taxon group k but the same group
k0
k

Level

sk

vk

vko

0
1
2
3
4
5
6
7
8
09
00
01
02

Species
Sub!genus
Genus
Tribe
Sub!family
Family
Super!family
Sub!order
Order
Series
Super!order
Sub!division
Class

82
78
61
56
48
30
28
22
03
8
6
5
1

0
1
2
3
4
5
6
7
8
09
00
01
02

0=2
5=8
7=8
01=4
10=3
11=8
16=3
33=3
43=8
50=3
54=5
74=2
099

abundance data for the calculation of D\ to

downweight the contributions of dominant species\
again identi_ed the Eastern Central and Southeastern
North Sea as taxonomically depauperate\ and gen!
erated similar values for all other coastal sectors "Fig[
2c#[ Using only presence:absence data\ the pattern in
taxonomic distinctness D between sectors was main!
tained for the Eastern Central and South!eastern
North Sea\ but the South!eastern Channel also had
low values of D "Fig[ 2d#[ Mean values of D for all
sectors except the Bristol Channel fell below that of
the theoretical mean\ indicating that most of the indi!
vidual sectors had lower average taxonomic range
than the entire survey area[
Comparison of taxonomic diversity D and taxo!
nomic distinctness D values determined from _sh
samples from all _shing station positions "Fig[ 3a#\
showed an underlying positive correlation\ but with
considerable variability in this relationship to the
lower right hand side of the _gure[ Here\ some samples
which were highly dominated and so had low values of
taxonomic diversity D\ represented assemblages which
were taxonomically broad "i[e[ high values of D#[
Note that the converse situation\ i[e[ taxonomically
restricted assemblages with high D\ cannot occur[ The
e}ect of data transformation on the calculation of D
was limited\ and the correlation between D calculated
using transformed and untransformed data remained
positive "Fig[ 3b#[ In contrast\ the apparently poor
correlation between D and D clearly showed the
in~uence of species abundances when calculating
taxonomic distinctness D "Fig[ 3c#[ Finally\ although
the use of a more re_ned weighting to re~ect the quan!
titative reduction in taxon richness on moving up the
hierarchy seems logical\ in fact the correlation between

Fig[ 2[ For the nine coastal sectors in Fig[ 0\ the taxonomic

diversity D "a# and the taxonomic distinctness D "b# of the
untransformed _sh abundance matrix\ and the taxonomic
distinctness D of square root transformed abundance data
"c#\ and the taxonomic distinctness D of presence:absence
data "d#[ For each sector the mean value with 84) con_dence
limits is shown[ The theoretical mean value of D calculated
by random sampling of species from the total species list is
marked on Fig[2d[

D calculated with and without the modi_ed weigh!

ting shows that they are highly correlated "Fig[ 3d#[
There was no relationship between the total number
of species in individual samples\ and their taxonomic
distinctness "D# calculated using presence:absence
data "Fig[ 4#[ Many of these D values\ however\ were
lower than the theoretical mean determined by ran!
dom sampling from the entire bottom!dwelling _sh
fauna\ a pattern already evident from Fig[ 2d[
Although most sample values fell within the 84) con!
_dence limit funnel\ a proportion had signi_cantly
lower values of D than the theoretical mean[
These values of D averaged over a quarter ICES
rectangle showed consistent trends in taxonomic dis!
tinctness from the Eastern Central North Sea to the

664
S[I[ Rogers\
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J[D[ Reynolds

Fig[3[ A comparison using all catch data between "a# taxonomic diversity D and taxonomic distinctness D\ "b# the root
transformed form of D against ordinary D\ "c# taxonomic distinctness D against ordinary D\ and "d# D using vko weights
against D using ordinary vk weights[

0888 British
Ecological Society
Journal of Animal
Ecology\ 57\ 658671

coastal waters west of the UK "Fig[ 5#[ The Eastern

Central and South!eastern North Sea\ and South!east!
ern Channel\ supported assemblages in which average
taxonomic distinctness was less than the theoretical
mean[ Assemblages close to\ and occasionally greater
than the theoretical mean\ occurred in the South!west!
ern North Sea\ along the south coast of England\ in
the Bristol Channel and in some parts of the western
Irish Sea "Fig[ 5#[
It is helpful when interpreting these regional di}er!
ences in taxonomic distinctness to identify which _sh
taxa have contributed most to patterns in D[ The
number of species belonging to each of the 03 orders
of bottom!dwelling _sh in the dataset were identi_ed
"Table 1#[ In the Eastern Central and South!eastern
North Sea\ which were the least taxonomically distinct
areas of all those surveyed\ most species belonged to
the orders Gadiformes "cod!like _shes#\ Perciformes
"bottom!dwelling or benthic _sh particularly drag!
onets of the family Callionymidae and gobies of the

family Gobiidae#\ Pleuronectiformes "~at_shes# and

Scorpaeniformes "gurnards\ Triglidae#[ These groups
contained almost all the species that were identi_ed
by Rogers et al[ "0888# as numerically most abundant
in the entire coastal demersal fauna\ and which typi!
_ed coastal sectors or discriminated between them
"Table 2#[ Only two elasmobranch orders were rep!
resented here\ and these only by a few species[ From
a total of 03 orders of _sh\ only 09 were represented
in the North Sea\ although the taxonomic distinctness
of the South!western North Sea was increased by the
presence there of greater numbers of more distantly
related cartilaginous species "Carcharhiniformes\
Rajiformes and Squaliformes# "Table 1#[ The bottom!
dwelling _sh fauna of the Channel was broadly similar
to that of the North Sea in terms of the relative dis!
tribution of species amongst phyla\ but the increased
abundance of cartilaginous species "Rajiformes# and
species from the order Perciformes\ together with
members of a more southerly fauna "e[g[ Zeiformes\

Zeus faber "L[## increased the taxonomic distinctness

of the region[ This pattern continued into the western
waters of the UK\ where additional representatives
of the southern fauna\ for example the electric ray
"Torpedinidae#\ mullet species "Mugiliformes#\ and
sun _sh "Tetradontiformes# were found[ The most
diverse fauna\ representing 03 orders of cartilaginous
and bony _shes\ was found in the Bristol Channel
"Table 1#[

665
Taxonomic
distinctness of _sh
communities

Discussion

Fig[ 4[ The departure from the theoretical mean taxonomic

distinctness D\ and 84) con_dence funnel\ of all individual
samples of bottom!dwelling _sh species calculated using
presence:absence data[ All values of D should fall within
the con_dence funnel assuming a null hypothesis that each
sample contains species randomly selected from the total
species list[

0888 British
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Journal of Animal
Ecology\ 57\ 658671

We have found clear spatial patterns in the taxonomic

diversity of bottom!dwelling _sh assemblages in the
coastal waters of NW Europe\ at both regional and
local scales[ The reduction in average taxonomic range
between the western waters of the UK and the sou!
thern North Sea mirrors similar patterns in the num!
ber of taxa represented\ and the relative abundance of
the species which comprise them "Fig[ 5^ Rogers et al[
0888#[ Although this suggests that taxonomic diversity
and traditional species diversity may indeed be posi!
tively correlated within this limited latitudinal range\

Fig[5[ Taxonomic distinctness D for samples grouped by quarter ICES rectangles[ Filled circles show areas with mean values
of D above the theoretical mean\ and open circles identify those regions with mean values below the theoretical mean[
Increasing bubble size re~ects the magnitude of departure from the mean[

666
S[I[ Rogers\
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J[D[ Reynolds

Table 1[ The mean number of species of demersal _sh that belong to the same order\ in each of the nine sectors sampled
Sector
Order

Rajiformes
Torpediniformes
Squaliformes
Carcharhiniformes
Anguilliformes
Gadiiformes
Lophiiformes
Gasterosteiformes
Perciformes
Pleuronectiformes
Scorpaeniformes
Zeiformes
Tetradontiformes
Mugiliformes

2[0
9[0
9[0
1[0
9[3
3[6
9[5
9[3
2[4
7[0
1[7
9[3
9[0
9[0

1[9
9[1
9
2[1
9[7
9[1
9
9[2
3[3
0[0
9
9
9
0[6

2[4
9[1
9
2[7
0[1
9[4
9
9[4
6[1
0[7
9
9
9
1[1

2[0
9[5
9
0[7
0[0
9[2
9
9[1
4[2
0[5
9[0
9
9
1[4

1[3
9
9
0[9
9[1
1[2
9
9[2
2[7
5[4
2[0
9[2
9
9

0[2
9
9
9[4
9
0[7
9
9[4
3[1
6[5
2[4
9[1
9
9

0[4
9
9[0
0[2
9[2
2[5
9
9[1
1[2
5[7
2[4
9
9
9

9[1
9
9
9[0
9[1
1[6
9
9[2
1[5
6[3
2[0
9
9
9

9[0
9
9
9[0
9[0
1[8
9[0
9[0
0[8
6[2
2[9
9
9
9

as suggested by Hall + Greenstreet "0887# for the

Northern North Sea\ the techniques used to calculate
these two types of index quantify di}erent aspects of
biodiversity[ This observation must therefore be a real
e}ect rather than an artefact of the methodology[
Records of samples which were taxonomically broad
but had highly dominated species abundances "i[e[ low
taxonomic diversity# "Fig[ 3a#\ highlight the additional
degree of resolution that can be achieved when cal!
culating these new indices on individual rather than
regionally aggregated samples[ It also shows that cor!
relations of the type described by Hall + Greenstreet
"0887# may only apply at relatively large geographical
scales[
These new indices incorporate information on taxo!
nomic relationships into diversity measures\ and also
support a statistical testing framework[ They are
therefore a valuable additional tool for interpreting
assemblage structure[ It is reasonable to assume that
the observed patterns in taxonomic diversity and dis!
tinctness are] "i# the inevitable consequence of the
regional patterns in natural processes\ or "ii# a result
of natural or man!made disturbance at a local scale\
or "iii# a combination of both[

NATURAL PROCESSES

0888 British
Ecological Society
Journal of Animal
Ecology\ 57\ 658671

Although not universally true\ many taxa show a clear

decrease in their species diversity from the tropics to
the poles "Pianka 0855^ Ekman 0856#[ One expla!
nation for this trend is that the tropics are larger in
area than temperate or tundra zones\ and experience
relatively high and uniform annual mean tem!
peratures "Rosenzweig 0881#[ In marine and fresh!
water systems\ the largest number of _sh species occur
in the tropics with progressively fewer towards polar
areas "Nelson 0883#[ While this rule can be applied
universally to groups that occupy the continental
shelf e[g[ coral reef _shes "Findley + Findley 0874#

and ~at_sh "Pauly 0883# there are some exceptions[

For example\ the deep water _sh fauna of the rise and
abyssal regions of the open oceans are more in~uenced
by the temperature di}erence throughout their depth
range than by the latitude\ and so for this group\
species diversity declines with increasing depth "White
0876#[ In the North!east Atlantic\ the warm water
fauna of the Mediterranean and Lusitanean regions
reaches its northern boundary on the continental shelf
to the west of the British Isles\ close to the English
Channel "Ekman 0856#[ In addition to the native _sh
fauna\ there are also representatives of the more sou!
therly Lusitanean fauna\ particularly those from the
order Perciformes\ such as the Sparidae\ Mullidae and
Labridae\ but also species from the families Zeidae\
Molidae\ and Mugillidae "Table 2#[ In most of the
North Sea\ there are two species groupings delimited
by the boundary between strati_ed and mixed water
masses approximately at the 39 m depth contour\ and
in much of the region\ gadoids "cod\ Gadus morhua
"L[#\ whiting\ Merlangius merlangus "L[# and pleuro!
nectids "dab\ Limanda limanda "L[## predominate
"Daan et al[ 0889#[
The role of local environmental disturbance and
productivity is particularly important in the southern
North Sea\ which\ unlike other regions that we exam!
ined\ is relatively shallow and supports an extensive
and homogeneous sandy substrate[ The benthic fauna
of this region is dominated by annelids\ molluscs and
crustaceans\ and it is maintained in an early stage of
succession partly by the in~uence of storm events and
tides[ In addition\ the nutrient!rich fresh water from
the Rivers Rhine and Elbe which ~ow into the German
Bight\ have created extensive eutrophication in
inshore waters which has contributed to these patterns
"Hickel\ Mangelsdorf + Berg 0882#[ The absence of
extensive reef!forming colonial invertebrates\ and the
varied fauna which can become associated with these
biogenic structures\ has reduced the range of niches

667
Taxonomic
distinctness of _sh
communities

Table 2[ The bottom!dwelling _sh taxa caught during the beam trawl surveys\ with their taxonomic hierarchy[ Those species
shown by Rogers et al[ "0887a# to comprise 9[3) or more of the total numerical abundance of the bottom!dwelling _sh catch
are marked by [ Amongst this group\ those which they additionally found to be consistent in typifying _sh communities
within sectors\ or consistent in discriminating between them\ are shown by 
CLASS

ORDER

FAMILY

Chondrichthys

Rajiformes

RAJIDAE

Torpediniformes

TORPEDINIDAE

Squaliformes
Carcharhiniformes

SQUALIDAE
SCYLIORHINIDAE
CARCHARINIDAE
TRIAKIDAE

Teleostei

Anguilliformes
Gadiformes

ANGUILLIDAE
CONGRIDAE
GADIDAE

PHYCIDAE

Lophiiformes
Gasterosteiformes

Perciformes

RANICIPITIDAE
MACROURIDAE
MERLUCCIIDAE
LOPHIIDAE
GASTEROSTEIDAE
SYNGNATHIDAE

CAPROIDAE
SPARIDAE
MULLIDAE
CEPOLIDAE
PERCICHTHYIDAE
LABRIDAE

ZOARCIDAE
PHOLIDAE
STICHAEIDAE
TRACHINIDAE
BLENNIIDAE
GOBIESOCIDAE
0888 British
Ecological Society
Journal of Animal
Ecology\ 57\ 658671

CALLIONYMIIDAE

blonde ray
thornback ray
small!eyed ray
spotted ray
starry ray
cuckoo ray
undulate ray
marbled electric ray
electric ray
spurdog
lesser spotted dog_sh
nurse hound
tope shark
smooth hound
starry smooth hound
eel
Conger eel
cod
haddock
whiting
poor cod
bib
pollack
common ling
_ve!bearded rockling
four!bearded rockling
shore rockling
three!bearded rockling
rockling species
lesser forkbeard
roundhead rat!tail
hake
angler
stickleback
greater pipe_sh
Nilsson|s pipe_sh
snake pipe_sh
sea horse
short snouted seahorse
boar!_sh
black sea!bream
red mullet
red band_sh
sea bass
bass family
cuckoo wrasse
ballan wrasse
rock cook
goldsinny
corkwing wrasse
eelpout
butter_sh
Yarrels blenny
snake blenny
greater weever
lesser weever
butter~y blenny
tompot blenny
two!spotted cling_sh
cling_sh family
dragonet family

Raja brachyura
Raja clavata
Raja microocellata
Raja montagui
Raja radiata
Raja naevus
Raja undulata
Torpedo marmorata
Torpedo nobiliana
Squalus acanthias
Scyliorhinus canicula
Scyliorhinus stellaris
Galeorhinus galeus
Mustelus mustelus
Mustelus asterias
Anguilla anguilla
Conger conger
Gadus morhua
Melanogrammus aegle_nus
Merlangius merlangus
Trisopterus minutus
Trisopterus luscus
Pollachius pollachius
Molva molva
Ciliata mustela
Enchelyopus cimbrius
Gaidropsarus mediterraneus
Gaidropsarus vulgaris
Gaidropsarus spp[
Raniceps raninus
Coryphaenoides rypestris
Merluccius merluccius
Lophius piscatorius
Gasterosteus aculeatus
Syngnathus acus
Syngnathus rostellatus
Entelurus aequoreus
Hippocampus ramulosus
Hippocampus hippocampus
Capros aper
Spondyliosoma cantharus
Mullus surmuletus
Cepola rubescens
Dicentrarchus labrax
Dicentrarchus spp[
Labrus mixtus
Labrus bergylta
Centrolabrus exoletus
Ctenolabrus rupestris
Crenilabrus melops
Zoarces viviparus
Pholis gunellus
Chirolophis ascanii
Lumpenus lampretaeformis
Trachinus draco
Echiichthys vipera
Blennius ocellaris
Blennius gattorugine
Diplecogaster bimaculata
Gobiesocidae
Callionymiidae

668
S[I[ Rogers\
K[R[ Clarke +
J[D[ Reynolds

Table 2[*continued[
CLASS

ORDER

FAMILY
GOBIIDAE

Pleuronectiformes

SOLEIDAE

PLEURONECTIDAE

SCOPHTHALMIDAE

BOTHIDAE
Scorpaeniformes

TRIGLIDAE

COTTIDAE
AGONIDAE
CYCLOPTERIDAE
LIPARIDAE
Zeiformes
Tetradontiformes
Mugiliformes

0888 British
Ecological Society
Journal of Animal
Ecology\ 57\ 658671

ZEIDAE
MOLIDAE
MUGILIDAE

rock goby
black goby
Stevens goby
Fries|s goby
goby family
sole
solenette
sand sole
thick back sole
plaice
~ounder
dab
lemon sole
witch
long rough dab
halibut
brill
turbot
megrim
Ekstroms topknot
Norwegian topknot
topknot
scald_sh
Imperial scald_sh
tub gurnard
red gurnard
grey gurnard
streaked gurnard
bull!rout
sea scorpion
hook!nose
lump sucker
sea snail
sea snail genus
John dory
sun_sh
thick lipped mullet
grey mullet family

Gobius paganellus
Gobius niger
Gobius gasteveni
Lesuerigobius friesii
Gobiidae
Solea solea
Buglossidium luteum
Pegusa lascaris
Microchirus variegatus
Pleuronectes platessa
Platichthys ~esus
Limanda limanda
Microstomus kitt
Glyptocephalus cynoglossus
Hippoglossoides platessoides
Hippoglossus hippoglossus
Scophthalmus rhombus
Scophthalmus maximus
Lepidorhombus whif_agonis
Phrynorhombus regius
Phrynorhombus norvegicus
Zeugopterus punctatus
Arnoglossus laterna
Arnoglossus imperialis
Trigla lucerna
Aspitrigla cuculus
Eutriglia gurnardus
Trigloporus lastoviza
Myoxocephalus scorpius
Taurulus bubalis
Agonus cataphractus
Cyclopterus lumpus
Liparis liparis
Liparis spp[
Zeus faber
Mola mola
Chelon labrosus
mugilidae

Fig[ 6[ Taxonomic distinctness D of the nine coastal sectors using presence:absence data "solid bars# "shown also in Fig[ 2d#\
showing the e}ect of removing all elasmobranch species "tinted bars#\ and all ~at_sh "pleuronectidae# "open bars# on the
calculation of the index[

679
Taxonomic
distinctness of _sh
communities

which the region can support[ This in turn has limited

the range of _sh species which can use these environ!
ments for food and shelter\ such as the herbivorous
mullets "Mugillidae# "Daan et al[ 0889#[

HUMAN IMPACTS

0888 British
Ecological Society
Journal of Animal
Ecology\ 57\ 658671

The North Sea has also experienced extensive manipu!

lation of its _sh communities by commercial _shing
activity\ and the extent of this disturbance\ and its
potential impact on the bottom!dwelling _sh assem!
blage\ deserves particular attention[ As one may
expect from a consistently high level of _shing activity\
the most obvious e}ects of the removal of _sh from
the higher trophic levels have been on the temporal
changes in size structure of communities "Pope 0878^
Greenstreet + Hall 0885^ Rice + Gislason 0885^ Rice
+ Kronlund 0886#[ The reduction in the abundance
of larger _sh "Pope + Knights 0871^ Rice + Gislason
0885^ Jennings\ Reynolds + Mills 0887# has caused
changes in the dominance of the smaller target and
non!target components of the assemblage\ as well as
the species composition of the catch[ Although several
studies undertaken in tropical waters show that locally
intense _shing activity can cause reductions in species
richness\ there are few examples of this e}ect in tem!
perate waters "Greenstreet + Hall 0885^ Rice + Gis!
lason 0885#^ and see review by "Jennings + Kaiser
0887#[ Is there any evidence that anthropogenic
activity has contributed to the lower taxonomic range
of populations in the southern North Sea\ compared
to elsewhere on the continental shelf<
There are some examples of commercial ground_sh
species in the North!east Atlantic disappearing from
large parts of their former range "Heessen + Daan
0885^ Rijnsdorp et al[ 0885^ Walker + Heessen 0885#[
Perhaps one of the best known involves the common
skate\ Raja batis "L[#\ "Brander 0870#[ The com!
bination of late age at maturity "00 years# large
maximum size "1 m# and a low potential rate of
population increase\ are thought to have been respon!
sible for its decline[ A similar decline has recently
been documented for the barndoor skate Raja laevis
"Mitchill# "Casey + Myers 0887#[ The presence of
these life!history characteristics was also directly
linked to decreasing trends in abundance of 07 inten!
sively exploited _sh stocks\ after accounting for
di}erences in their _shing mortality "Jennings et al[
0887#[ The elasmobranchs have some or all of these
characteristics\ and dog_sh\ and several species of ray
"Squaliformes and Carcharhiniformes#\ which gen!
erally have a widespread distribution throughout the
North!east Atlantic\ are exploited by commercial
_sheries "Holden 0863^ Quero + Cendrero 0885^ ICES
0886^ Walker\ Howlett + Millner 0886#[ In the Bay of
Biscay\ the long!term decline in abundance of three
species of ray "Raja batis "L[#\ Raja brachyura "Lafont#
and Raja clavata "L[##\ and other species of large
bodied carcharhiniform sharks\ was attributed to the

long!term e}ects of bottom trawling throughout the

area "Quero + Cendrero 0885#[ Of all the families
which comprise the total _sh assemblage therefore the
elasmobranchs are the most likely to show e}ects of
commercial activity\ if it is present[
To what extent do the elasmobranchs in~uence
the spatial patterns in taxonomic distinctness that we
identi_ed in Fig[ 5< To investigate this further\ values
of D were recalculated for each sector\ but using a
revised dataset in which all elasmobranchs\ i[e[ mem!
bers of the Chondrichthyes "Table 2## were assumed
to be absent from catches[ The result of this analysis
describes the taxonomic distinctness of the bottom!
dwelling\ non!elasmobranch _shes\ and should
exclude the potentially confounding e}ects of human
activity on this group to reveal the underlying bio!
geographic pattern[ Di}erences in mean D between
sectors using this revised dataset were less signi_cant
"F statistic  7=3# than when previously calculated
using the full dataset "F  31=8# "Fig[ 6#[ In particular\
mean taxonomic distinctness in all sectors of the
North Sea appeared only marginally less than those
west of the UK "Fig[ 6#[ This is not simply a scale
e}ect in which a reduced dataset leads to reduced
discrimination[ When the calculations were repeated
but with all the ~at_sh "Pleuronectiformes\ a similar!
sized group# excluded from the dataset instead of the
elasmobranchs\ the e}ect was barely detectable "Fig[
6#[ Mean values of D retained the same general
relationships between sectors\ and the di}erences
between sector means were still relatively large
"F  20[6#[ This analysis con_rms that the elasmo!
branchs have a major in~uence on the taxonomic dis!
tinctness of the North!east Atlantic fauna\ and that
when they are arti_cally excluded from the samples\
the remaining species have a more uniform taxonomic
range throughout the region[ In such a widely dis!
tributed bottom!dwelling group\ this e}ect is surpris!
ing[ One possible explanation is that the adverse
e}ects of high _shing mortality have in~uenced the
distribution and abundance of some elasmobranch
species to create an arti_cial pattern[ Alternatively\ it
may result from di}erent levels of _shing activity
within coastal waters[ Further analysis of time!series
changes in the distribution and abundance of elasmo!
branchs\ their life!history characteristics and their pat!
terns of exploitation\ will be necessary to con_rm the
extent of the role of high _shing e}ort in the decline
of some of these species[
In the event of perturbations caused by either
human impact or changing environmental conditions\
Tilman "0885# claims that the taxonomic range of
an assemblage will be important for maintaining the
stability of the ecosystem[ In some marine invertebrate
communities there is considerable redundancy at the
species level\ such that similar patterns in structure
can be identi_ed at a higher taxonomic level[ This
suggests that there may also be several species which
perform the same functional role "Ferraro + Cole

670
S[I[ Rogers\
K[R[ Clarke +
J[D[ Reynolds

0881^ Clarke + Warwick 0887a#[ Marine _sh com!

munities with low levels of species redundancy and
where few species ful_l key functional roles\ for exam!
ple some coral reef ecosystems\ may be least able to
withstand environmental change "Hughes 0883^ Jen!
nings + Kaiser 0887#[ These indices of taxonomic
diversity and distinctness can therefore provide valu!
able information on the extent to which _sh assem!
blages are able to resist change\ and this will become
increasingly relevant in our interpretation of changes
in assemblage structure[

Acknowledgements
The authors wish to thank Uli Damm "Germany#\
Adriaan Rijnsdorp "The Netherlands#\ Willy Vanhee
"Belgium#\ and their colleagues\ for making these
international data available for this analysis[ Nicholas
Dulvy and Suzanne Mills kindly helped to compile
the taxonomy[ This work was supported by MOU"A#
of the Ministry of Agriculture\ Fisheries and Food\
and the participation of K[R[C[ was funded jointly by
MAFF "AE0002# and the NERC Marine Biodiversity
programme at CCMS Plymouth Marine Laboratory[
We thank Simon Greenstreet and an anonymous ref!
eree for helpful comments[

References

0888 British
Ecological Society
Journal of Animal
Ecology\ 57\ 658671

Baltz\ D[M[ "0880# Introduced _shes in marine systems and

inland seas[ Biological Conservation\ 45\ 040066[
Brander\ K[ "0870# Disappearance of common skate Raia
batis from the Irish Sea[ Nature\ 189\ 3738[
Casey\ J[M[ + Myers\ R[A[ "0887# Near extinction of a large\
widely distributed _sh[ Science\ 170\ 589581[
Clarke\ K[R[ + Warwick\ R[M[ "0887a# Quantifying struc!
tural redundancy in ecological communities[ Oecologia\
002\ 167178[
Clarke\ K[R[ + Warwick\ R[M[ "0887b# A taxonomic dis!
tinctness index and its statistical properties[ Journal of
Applied Ecology\ 24\ 412420[
Connell\ J[H[ "0867# Diversity in tropical rain forests[
Science\ 088\ 02910209[
Daan\ N[\ Bromley\ P[J[\ Hislop\ J[R[G[ + Nielsen\ N[A[
"0889# Ecology of North Sea _sh[ Netherlands Journal of
Sea Research\ 15\ 232275[
Ekman\ S[ "0856# Zoogeography of the Sea[ Sidgwick and
Jackson\ London
Ferraro\ S[P[ + Cole\ F[A[ "0881# Taxonomic level su.cient
for assessing a moderate impact on macrobenthic com!
munities in Puget Sound\ Washington\ USA[ Canadian
Journal of Fisheries and Aquatic Science\ 38\ 00730077[
Findley\ J[S[ + Findley\ M[T[ "0874# A search for pattern in
butter~y _sh communities[ American Naturalist\ 015\ 799
705[
Gee\ J[M[ + Warwick\ R[M[ "0885# A study of global bio!
diversity patterns in the marine motile fauna of hard sub!
strata[ Journal of the Marine Biological Association\ UK \
65\ 066073[
Greenstreet\ S[P[R[ + Hall\ S[J[ "0885# Fishing and the
ground!_sh assemblage structure in the north!western
North Sea] an analysis of long!term and spatial trends[
Journal of Animal Ecology\ 54\ 466487[
Hall\ S[J[ + Greenstreet\ S[P[R[ "0887# Taxonomic dis!

tinctness and diversity measures] responses in marine _sh

communities[ Marine Ecology Progress in Series\ 055\ 116
118[
Heessen\ H[J[L[ + Daan\ N[ "0885# Long!term trends in ten
non!target North Sea _sh species[ ICES Journal of Marine
Science\ 42\ 09590967[
Hickel\ W[\ Mangelsdorf\ P[ + Berg\ J[ "0882# The human
impact in the German Bight] eutrophication during three
decades "085180#[ Helgolander Meeresunters\ 36\ 132
152[
Holden\ M[J[ "0863# Problems in the rational exploitation of
elasmobranch populations and some suggested solutions[
Sea Fisheries Research "ed[ F[R[ Harden Jones#\ pp[ 006
026[ Elek Science\ London[
Hughes\ T[P[ "0883# Catastrophes\ phase shifts and large!
scale degradation of a Caribbean reef[ Science\ 154\ 0436
0440[ICES[ "0886# Report of the Study Group on Elasmo!
branch Fishes ICES CM 0886:G] 1[
Jennings\ S[ + Kaiser\ M[J[ "0887# The e}ects of _shing on
marine ecosystems[ Advances in Marine Biology\ 23\ 190
241[
Jennings\ S[\ Reynolds\ J[D[ + Mills\ S[D[ "0887# Life history
correlates of responses to _sheries exploitation[ Pro!
ceedings of the Royal Society of\ London\ 154\ 222228[
Ludwig\ J[A[ + Reynolds\ J[F[ "0877# Statistical Ecology] a
Primer on Methods and Computing[ John Wiley and Sons\
New York[
Magurran\ A[E[ "0877# Ecological Diversity and its Measure!
ment[ Chapman + Hall[ London[
McEachran\ J[D[ + Miyake\ T[ "0889# Phylogenetic inter!
relationships of skates] a working hypothesis "Chon!
drichthyes\ Rajoidei#[ Elasmobranchs as Living Resources]
Advances in the Biology\ Ecology\ Systematics\ and the
Status of the Fisheries "eds H[L[ Pratt\ S[H[ Gruber + T[
Taniuchi#\ pp[ 174293[ NOAA Technical Report NMFS
89[
McEachran\ J[D[\ Dunn\ K[A[ + Miyake\ T[ "0885# Inter!
relationships of the Batoid _shes "Chondrichthyes] Bato!
idea#[ Interrelationships of Fishes "eds M[L[J[ Stiassny\
L[R[ Parenti + G[D[ Johnston#\ pp[ 5273[ Academic
Press\ San Diego[
Nelson\ J[S[ "0883# Fishes of the World\ 2rd edn[ John Wiley
+ Sons\ New York[
Pauly\ D[ "0883# A framework for latitudinal comparisons
of ~at_sh recruitment[ Netherlands Journal of Sea
Research\ 21\ 096007[
Pearson\ T[H[ + Rosenberg\ R[ "0867# Macrobenthic suc!
cession in relation to organic enrichment and pollution of
the marine environment[ Oceanography and Marine
Biology\ Annual Review\ 05\ 118200[
Petraitis\ P[S[\ Latham\ R[E[ + Niesenbaum\ R[A[ "0878#
The maintenance of species diversity by disturbance[ Quar!
terly Review of Biology\ 53\ 282307[
Pianka\ E[R[ "0855# Latitudinal gradients in species diversity]
a review of concepts[ American Naturalist\ 099\ 2235[
Pope\ J[G[ + Knights\ B[J[ "0871# Comparison of the length
distributions of combined catches of all demersal _shes in
surveys in the North Sea and at Faroe Bank[ Canadian
Special Publication of Fisheries and Aquatic Sciences\ Vol
48 "ed[ M[C[ Mercer#\ pp[ 005007[ Government of
Canada\ Ottawa[
Pope\ J[G[ "0878# Fisheries research and management for the
North Sea] the next hundred years[ Dana\ 7\ 2232[
Quero\ J[!C[ + Cendrero\ O[ "0885# Incidence de la peche
sur la biodiversite ichtyologique marine] Le bassin d|Ar!
cachon et le plateau continental sud Gascogne[ Cybium\
19\ 212245[
Rice\ J[ + Gislason\ H[ "0885# Patterns of change in the
size spectra of numbers and diversity of the North Sea
assemblage\ as re~ected in surveys and models[ ICES Jour!
nal of Marine Science\ 42\ 01030114[

671
Taxonomic
distinctness of _sh
communities

0888 British
Ecological Society
Journal of Animal
Ecology\ 57\ 658671

Rice\ J[C[ + Kronlund\ A[R[ "0886# Community analysis

and ~at_sh] diagnostic patterns\ processes\ and inference[
Journal of Sea Research\ 26\ 290219[
Rijnsdorp\ A[D[\ Van Leewen\ P[I[\ Daan\ N[ + Heessen\
H[J[L[ "0885# Changes in abundance of demersal _sh spec!
ies in the North Sea between 0895 and 0898 and 088984[
ICES Journal of Marine Science\ 42\ 09430951[
Rogers\ S[I[\ Maxwell\ D[\ Rijnsdorp\ A[D[\ Damm\ U[ +
Vanhee\ W[ "0888# Fishing e}ects in northeast Atlantic
Shelf Seas] patterns in _shing e}ort\ diversity and com!
munity structure[ IV[ Can comparisons of species diversity
be used to assess human impacts on coastal demersal _sh
faunas in the Northeast Atlantic< Fisheries Research\ 39\
024041[
Rogers\ S[I[\ Rijnsdorp\ A[D[\ Damm\ U[ + Vanhee\ W[
"0887# Demersal _sh populations in the coastal waters of
the UK and continental NW Europe from beam trawl
survey data collected from 0889 to 0884[ Journal of Sea
Research\ 26\ 68091[
Rosenzweig\ M[L[ "0881# Species diversity gradients] we
know more and less than we thought[ Journal of Mammo!
logy\ 62\ 604629[
Rosenzweig\ M[L[ "0884# Species Diversity in Space and
Time[ Cambridge University Press\ Cambridge[
Somer_eld\ P[J[\ Olsgard\ F[ + Carr\ K[R[ "0886# A further
examination of two new taxonomic distinctness measures[
Marine Ecology Progress in Series\ 043\ 292295[
Sousa\ W[P[ "0868# Disturbance in marine intertidal boulder

_elds] the nonequilibrium maintenance of species diversity[

Ecology\ 59\ 01140128[
Tilman\ D[ "0885# Biodiversity] population versus ecosystem
stability[ Ecology\ 66\ 249252[
Walker\ P[\ Howlett\ G[ + Millner\ R[ "0886# Distribution\
movement and stock structure of three ray species in the
North Sea and eastern Channel[ ICES Journal of Marine
Science\ 43\ 686797[
Walker\ P[A[ + Heessen\ H[J[L[ "0885# Long!term changes
in ray populations in the North Sea[ ICES Journal of
Marine Science\ 42\ 09740982[
Warwick\ R[M[ + Clarke\ K[R[ "0882# Comparing the sever!
ity of disturbance] a meta!analysis of marine macrobenthic
community data[ Marine Ecology Progress in Series\ 81\
110120[
Warwick\ R[M[ + Clarke\ K[R[ "0884# New {biodiversity|
measures reveal a decrease in taxonomic distinctness with
increasing stress[ Marine Ecology Progress in Series\ 018\
290294[
Warwick\ R[M[ + Clarke\ K[R[ "0887# Taxonomic dis!
tinctness and environmental assessment[ Journal of
Applied Ecology\ 24\ 421432[
White\ B[N[ "0876# Oceanic anoxic events and allopatric
speciation in the deep sea[ Biological Oceanography\ 4\
132148[
Received 11 June 0887^ revision received 3 November 0887