Freshwater Biology (1981) 11, 99-120

A comparative study of seven grabs used for sampling

benthic macroinvertebrates in rivers

J. M. ELLIOTT and C. M. DRAKE Freshwater Biological Association, Windermere Laboratory.

England

SUMMARY. After considering the large number of grabs described in the

literature, seven grabs of weight < 25 kg were chosen for manual operation
from a small boat: Van-Veen grab, weighted and unweighted Ponar grabs,
Friedinger version of the Petersen grab, Dietz-La Fond mud-snapper,
pole-operated Birge-Ekman grab and pole-operated Allan grab.
Random samples (number of sampling unitsn = 10) were taken in a large
tank with a known number of 2-mm cylindrical plastic pellets amongst stones
of uniform size. Separate experiments were performed with four sizes of
stones (model ranges: 2-4 mm, 8-16 mm, 16-32 mm, 32-64 mm). Stratified
random samples {n = 10) were taken in rivers and the modal particle sizes at
four sites were 0.004-0.06 mm, 0.5-2 mm, 16-64 mm and 64-128 mm. All
grabs usually took a representative sample of the substratum at each site
with no strong bias towards a particular particle size. The general perfor-
mance of the Friedinger, Dietz-La Fond and Allan grabs was poor, except
on a muddy bottom, with frequent failure to operate, small samples of
substratum and a mean depth of penetration < 3 cm in all substrata except
mud for the Dietz-La Fond and Allan grabs. The Van-Veen and Birge-
Ekman grabs sampled to a mean depth < 3 cm in mud and fine gravel (2-4
mm), but the Birge-Ekman jammed frequently in fine gravel. Both Ponar
grabs operated well and sampled to a mean depth 3^5 cm in mud and fine
gravel, > 3 cm when small stones (8-16 mm) were present and 2 cm
(weighted Ponar only) when larger stones (> 16 mm) were present in a
gravel bottom. The mean depth was <0.8 cm for all grabs when larger stones
(>16 mm) were predominant on the bottom.
In the tank experiments with pellets, the efficiencies for the total catches
of the Friedinger, Dietz-La Fond and Allan grabs were low with values
<45% for fine gravel (2-4 mm), < 22% for small stones (8-16 mm) and
<5% for a substratum of larger stones (>16 mm). If 50% is the minimum
acceptable efficiency, then the Ponar, Van-Veen and Birge-Ekman grabs
were adequate for fine gravel, only the two Ponar grabs were adequate for
small stones and no grabs were adequate for sampling a substratum of larger
stones (>16 mm).

Correspondence: J. M, Elliott. Freshwater Biologi-

cal Association, The Ferry House, Ambleside, Cum-
bria LA22 OLP. England.

0046-5070/81/0400-0099 $02.00 © 1981 Blackwel! Scientific Publications

100 J. M. Elliott and C. M. Drake

In field trials, the relative abundances of major taxa were similar for most
grabs at each site; Friedinger and Dietz-La Fond grabs were the major
exceptions. In terms of both mean number of taxa and mean number of
invertebrates m"^ the Ponar, Birge-Ekman and Allan grabs performed well
on the predominantly muddy substratum at site 1, but only the weighted
Ponar grab performed adequately on the predominantly gravel bottom with
some large stones (>16 mm) at site 2. All grabs performed badly when
larger stones (>16 mm) were predominant on the bottom (sites 3, 4).
The relationship between the variances and means of the samples taken
with each grab followed a power law for the catches of pellets in tank
experiments, and for major taxa and total numbers at each site in field trials.
Values of exponents in the power law lay within the range 1.14-2.34. The
coefficient of variation was also frequently related to the sample mean and
was an unreliable statistic for comparing the precision of grabs.

Introduction limited to two or three grabs, e.g., for marine

Macroinvertebrates are frequently used to
assess water quality in rivers and are usually
defined as invertebrates that are retained by a
net or sieve with an aperture of 0.6 mm (Weber,
1973). A smaller aperture of 0.25 mm may be
necessary for special purposes, e.g., to ensure
the capture of early instars. Samplers for mac-
roinvertebrates can be divided into three broad
categories: traps, colonization samplers and
immediate samplers. The advantages and disad-
vantages of these three types are summarized by
Elliott, Drake & Tullett (1980). who also con-
sider the general problem of choosing a sampler
and emphasize that the final choice is usually
determined by the objectives of the investiga-
tion. Immediate samplers are used for the rapid
removal of benthic macroinvertebrates from the
substratum and a wide variety of devices have
been invented for scraping, digging, coring or
sucking samples from the bottom. The literature
on these samplers is listed in an annotated bib-
liography that includes not only freshwater sam-
plers but also marine samplers that have been, or
could be. used in fresh water (Elliott & Tullett,
1978). As this review of the literature showed
that grabs are frequently used for sampling in
deep water (depth >1 m), several grabs were
selected and tested in a series of trials in both the
field and the laboratory. The results of this work
are described in the present paper.
Several workers have compared the perfor-
mances of grabs but these studies are usually
benthos: Petersen and Van-Veen (Thamdrup,
1938; Ursin, 1954; Birkett, 1958; Kutty &
Desai. 1968); Smith-Mclntyre and Van-Veen
(Wigley, 1967; Bhaud & Duchene, 1977);
Petersen, Smith-Mclntyre and Van-Veen (Gal-
lardo, 1965); for freshwater benthos: Birge-
Ekman and Ponar (Howmiller, 1971); Birge-
Ekman, Ponar and orange-peel (Hudson,
1970); Birge-Ekman, Ponar and Petersen
(Weber. 1973). Comparisons of more than
three grabs are limited to one marine study with
five grabs: Petersen, Van-Veen, orange-peel,
Smith-Mclntyre, Holme double scoop (Reys,
1964); and two studies on lake benthos with six
grabs: Birge-Ekman, Ponar, Shipek, Dietz-La
Fond and Franklin-Anderson in both studies
with Petersen in one (Sly, 1969) and tall Birge-
Ekman in the other (Flannagan, 1970). Seven
grabs were used in the present study and trials
were performed in a test-tank with plastic pellets
and in rivers with macro-invertebrates amongst
stony substrata.
Selection and description of grabs
Grabs are samplers with jaws that are forced
shut by weights, lever-arms, springs or cords.
Over sixty grabs have been described with about
equal numbers from marine and freshwater
studies (Elliott & Tullett, 1978). Most of these
grabs are simply modifications of five basic
types: Petersen, Van-Veen, Birge-Ekman,

Smith-Mclntyre and orange-peel. TTie larger
grabs are very heavy and require a large boat
with a power-winch, e.g., box-corer grab at 750
kg (Reineck. 1963). Campbell grab at 410 kg
(Hartman, 1955), Okean grab at 150 kg (Lisit-
syn & Udintsev, 1955), pneumatic version of
Birge-Ekman at 130 kg (Murray & Charles,
1975), single-scoop at UO kg and double-scoop
at 115 kg (Holme, 1949. 1953), orange-peel
grab at 86 kg (Reys, 1964; Holme & Mclntyre,
1971), Shipek scoop at 70 kg (Hydro Products,
1973) and Smith-Mclntyre grab at 45 kg (Smith
& Mclntyre, 1954). All these grabs were
rejected because one requirement of the present
study was that the grabs had to be operated
manually from a small boat and therefore had to
weigh less than 25 kg. Lighter versions of the
orange-peel grab, weighing about 20-25 kg,
were considered but were rejected because it
was thought unlikely that the four jaws would
close completely on a stony substratum (see also
the criticisms of Thorson, 1957) and because
there is a marked decrease in the sampling area
from the surface of the substratum to the max-
imum penetration of the jaws (Hudson, 1970).
After considering the various specifications of
grabs, especially their weights, the following
seven versions were chosen (Fig. 1): Van-Veen
(Van- Veen, 1936; Thamdrup, 1938; Thorson,
1957), weighted and unweighted Ponar-type
(Powers & Robertson, 1967), Friedinger
(Naumann. 1925), Dietz-La Fond (La Fond &
Dietz, 1948), Birge-Ekman (Ekman, 1911;
Birge, 1921) and Allan (Allan, 1952).
The Van-Veen grab weighs 21 kg, has a sam-
pling area of 0.106 m^ and has jaws that are
closed by a pincer-like action of two long arms
(Fig. la). Both Ponar grabs have a sampling area
of 0.056 m^ and jaws that are closed by the
scissor-like action of two pivoted arms; the
unweighted and weighted versions weigh 14 kg
and 23 kg, respectively (Fig. Ib.c). The
Friedinger grab is a small version of the Petersen
grab and weighs 8 kg, has a sampling area of
0.030 m^ and has jaws that are closed by wires
that cross from one jaw to the other (Fig. Id).
The Dietz-La Fond grab is one of the largest and
heaviest of the 'mud-snappers' that are foot-
triggered grabs with the jaws closed by a power-
ful spring (Fig. le). This grab weighs 21 kg and
has a sampling area of only 0.016 m^. A pole-
operated version of the Birge-Ekman grab was
chosen and can be used in water up to 3 m deep
Comparative study of grabs 101
(Fig. If). The weight is only 7 kg and the sampl-
ing area is 0.023 m^ A second pole-operated
grab, the Allan grab, was also chosen and can be
used to a depth of 3.3 m(Fig. lg).Theweight is8
kg and the sampling area is 0.035 m'. Whilst the
jaws of the Birge-Ekman grab are closed by
strong springs, those of an Allan grab are closed
by a manually-operated rod and levers.
Methods
Laboratory experiments
Tliese were performed in a large tank (length
of sides 103 cm, height 165 cm) made of glass-
reinforced polyester resin. The depth of water in
the tank was maintained in the range 1.3-1.4 m.
A wooden tray (length of sides 73.5 cm, internal
height 22.5 cm) was filled with stones of similar
size to form a uniform substratum with a depth
of 8-10 cm. Small cylindrical pellets of plastic
(modal length and diiimeter 2 mm) represented
'invertebrates' and these were scattered over the
tray in two layers; one layer of white pellets at a
depth of about 3 cm and one layer of blue pellets
just below the surface of the substratum. The
pellets were chosen because their size and
specific gravity were similar to those of benthic
macroinvertebrates. A constant weight of pel-
lets was used for each experiment and a prelimi-
nary study showed that the mean density (±95%
CL) of pellets in the tray was 12500 ± 354 with
6330 ± 225 white pellets and 6170 ± 117 blue
pallets. These values are equivalent to a density
of 23142 ± 655 m-^ with 11715 ± 416 m"^ for
the pellets at a depth of 3 cm and 11427 ± 217
m"- for the pellets near the surface. Therefore,
thedensity of pellets in the tray was high and Just
below the highest densities attained in the field
(cf. with values In Fig. 6). In one series of exper-
iments with an unweighted Ponar grab, the den-
sities of white and blue pellets were reduced to
80%, 60%, 40% and 20% of their usual values
of 6330 and 6170, respectively.
Four sizes of stones were used in the experi-
ments. Fine gravel was the smallest with a modal
size range of 2-4 mm and an overall range of
0.1-5 mm. As the larger stones were not round,
the longest axis was measured (standard analysis
of Pettijohn, 1957) for a random sample of 300
stones. The modal size ranges were 8-16 mm
102 J. M. Elliott and C. M. Drake
Open
(d)
Open
FIG. 1. The seven grabs used in this study (the vertical line next to each grab is equivalent to 10 cm), (a) Van-Veen;
(b) weighted Ponar; (c) Ponar; (d) Friedinger; (e) Dietz-La Fond; (f) Birge-Ekman; (g) Allan.
(overall range 3-16 mtn), 16-32 mm (overall
range 8-35 mm) and 32-64 mm (overall range
10-66 mm).
One stone size was used in each series of
experiments and a sample often replicate sampl-
ing units was taken with each grab. The number
of replicates per tray varied from three with the
large Van-Veen grab to eight with the small
Dictz-La Fond grab. Therefore, the trays had to
be changed frequently for the larger grabs. The
pellets were separated from the stones by flota-
tion in carbon tetrachloride and the white and
blue pellets in each sampling unit were counted
Open Closed
tg)
separately. An estimate of the gross volume of
the stones from each sampling unit was made by
shaking down the stones under water in a
measuring cylinder so that the gross volume was
the volume of the stones plus the volume of the
spaces between the stones. This measure, rather
than the net volume, was thought to be closer to
the capacity of each grab.
Field trials
Samples were taken at four sites and the modal
size of the substratum increased progressively
 Comparative study of grabs 103

from site I to site 4. Two rivers in the English Sphaerium, Hydra, small Baetis. After the
Lake District were used for the field trials; Black invertebrates had been removed, the gross vol-
Beck in August 1978 (site 1) and the River ume of the stones from each sampling unit was
Rothay in June 1979 (sites 2, 3) and August measured for samples taken from the Rothay.
1978 (site 4). Black Beck is the main inflow of As most of the mud had been removed from the
Esthwaite Water and site 1 was situated about Black Beck samples in the field, the volume of
200 m from the mouth of the river. Water depth substratum in each sampling unit could not be
at the site was 1-2.5 m with a width of 3-5 m and measured in the laboratory. Therefore, a second
a flow of 10—40 cm s"'. The bottom is predomin- series of samples was taken from Black Beck in
antly mud (particle size 0.004-0.06 mm) with October 1979 and these samples were used to
some fine gravel (particle size 0.06-5 mm), determine the volumes of substratum.
numerous packets of leaves and broken twigs, All the statistical methods used to analyse the
and small patches of macrophytes. e.g., Callit- data are described in detail by Elliott (1977). A
riche sp., Elodea canadensis Michx. and Spar- 5% level of significance was used in all statistical
ganium erectum L. The River Rothay is the main tests.
inflow for Windermere and sites 2-4 were situ-
ated about 350 m from the mouth of the river.
Water depth at each site was 1-2 m with a width Results
of 5-8 m and a flow of 10-50 cm s ' . The
bottom is stony with little mud, no macrophyles General performance characteristics of the grabs
and sparse clumps of moss on larger stones.
Samples in June 1979 were taken at two sites, Bite profiles were determined by cutting ten
each with a bottom of loose stones and a modal profiles with each grab in wet sand (modal size
size range of 0.5-2 mm (overall range <1-I28 range 0.25-0.50 mm) and also in fine gravel
mm) at site 2 and 16-64 mm (overall range (modal size range 2-4 mm) in trays on land. The
<1-128 mm) at site 3. Samples in August 1978
were taken at site 4 which has a 'cemented'
bottom of tightly packed stones with a modal
size range of64-l28 mm (overall range < 1-128 (a ! Van-Veen
mm). These size ranges were obtained from a
dry weight analysis of particle sizes in all samples ( b} Weighted Ponar
taken by the grabs (method described in detail
by Welch, 1948). Similar results were obtained
for random samples taken from the bottom with C ) Poncjr
a pond net.
At each site, a stretch of river, 25 m long, was
Id ) FriedJnger
divided into five sections, each 5 m long. Two
sampling units were taken at random from each
5-m section to give a stratified - random sam-
e ) Dieli-LoFoncJ
ple of ten sampling units for each grab. If a grab
did not catch anything after three attempts, a
value of zero was recorded for the sampling unit.
Each catch was emptied into a bath and then
concentrated by filtration through a nylon bag
(aperture 0.595 mm). The amount of mud that (g) Allan
passed through the bag was negligible in the
Rothay samples but considerable in the Black
Beck samples. All catches were preserved in 10 20 30 40
10% formalin and hand sorted in the laboratory. Width Icm5
Most taxa were identified to species, but major
exceptions were oligochaetes identified to FIG. 2. Typical profiles cut by the seven grabs in fine
gravel; (a) Van-Veen; (b) weighted Ponar; (c) Ponar;
families, chironomids to sub-families or tribes (d) Friedinger; (e) Dietz-La Fond; (Q Birge-Ekman;
and the following to genera: Folyceiis, Pisidium, (g) Allan.
iO4 J. M. Elliott and C. M. Drake

Site Z Site 3
4000 p i-i 4000
0 3000 a a
m
2000 - 2000 2000

4000
o
-

-r-n-H 1 . r
2000 -

IZOOOr Totol

6000

0
OS
I 2 4 8 32 IZ8
le 64
Q: . 0
0-5
I 2 4 e 32 128
16 £4
. 8 32 128
16 64
Particia »\tt (mm)

FIG, 3. Weight of substratum in each particle-size class for samples taken by each grab at sites 2-4; (a) Van-Veen,
(b) weighted Ponar. (c) Ponar, (d) Friedinger, (e) Dietz-La Fond, (f) Birge-Ekman, (g) Allan.
 Comparative study of grabs 105

profiles were very similar in the two substrata

and were virtually the same for the ten replicates
with each grab, but varied considerably between
the seven grabs (Fig. 2). Profiles for the Van-
Veen, Ponar and Friedinger grabs were similar • « •
o <s (O
00 r-.
to the U-shaped profiles described for Van- +1 1 +1 +1 +1 (N
Veen and Petersen grabs by Birkett (1958), u 1 O u-l +1
.t: 4 o ro •t ON ro
whilst the profiles for the Dietz-La Fond and
Birge-Ekman grabs were closer to the
inverted-V profiles obtained for Van-Veen and
Petersen grabs by Gallardo (1965). Small ridges +1 +1 +1 +1 +1 +1 +1
were left in the centre of the profiles eut by the •T c
00 r-
tNCTvr^ 00 \o
00 - - r-- (N o\
Van-Veen and especially the Dietz-La Fond and
Birge-Ekman grabs. Similar ridges were 00 (N
observed by Gallardo (1965). Only the Allan \Ci O O O ~^ ^ ^^

grab produced an almost rectangular profile, +1 +1 +1 +1 +1 +1 +1

'^ C^ F^ "^ C^ O^ C^
and there was a marked reduction in Ihe sampl- O '—< »/^ ^H ^H
ing area with depth for all the other grabs.
O 00 ^
The substratum in each series of tank experi- f*^ O fN u^ ^^ »-^ ^
ments and in the field trials at site 1 was fairly
+1 +1 +1 +1 +1 +1 +1
uniform, but there was considerable variation in 00 iri rs «-i oo iO M*i
<o v i rN - - r-4 «*i (S
particle size at sites 2, 3 and 4. Weight distribu-
tions of particle sizes at each of these sites were
remarkably similar for most grabs (Fig. 3) and irj lO U-) ^ - in
were also similar to the weight distribution of •<t ao ^^
ro (N ^ r-
U-) o »O
vO <M «
particle sizes for the total substrata representa- +1 +1 +t +1 W -H +1
— O- O rs u-l "It •*
tive of the bottom conditions at each site (see
methods). Exceptions were the Allan grab at site
2 and the Friedinger and Dietz-La Fond grabs at
site 3, these grabs showing a slight bias towards
the larger stones. As the weights of substrata +1 +1 +1 +1 +1 +1 +1

taken in these two samples were very low, little 'O OO O 00 V l V^

oo r o 00 ro " ^
CTN

importance can be attached to these discrepan- (O

cies and it can be generally concluded that the r) lo O
ro C ^ ^G ^ r*^ ^
--• •©
•—' r O V I »—< 0 0 *—< CT^
grabs usually took a representative sample of the +1 +1 +1 +1 +1 +1 tt
substratum at each site. vo t^ in -t — —
O r^ v^ S m ro CT>
Both the weight (Fig. 3) and volume (Table 1) ro ro " ro ro

of substrata in the samples varied considerably

between grabs and sites. These variations were £. S r—1 —
o ^n
due to the different sampling areas of the grabs — «
+1 +1 +1 +1 +1 +1 +1
00 —

and variations in substratum particle size. As the •<t — O\ 00 -O

(N ro -O
latter increased, the amount of substratum taken So ro 00 r--

by each grab usually decreased significantly in

both the tank and field experiments. Major "S
exceptions were the Dietz-La Fond grab with no
significant decrease in the tank experiments and
between sites 3 and 4 in the field trials, and the
rather erratic changes in the volumes taken by
the Birge-Ekman and Allan grabs. There were
no obvious reasons for these departures from the ,2-H
general pattern but, as will be shown later, there
were no corresponding changes in the catches of
plastic pellets and these showed a marked dec-
106 J. M. Elliott and C. M. Drake

rease with increasing substratum size for all

grabs. Although the sampling areas of the grabs
must have affected the amounts of substrata "-•
fN
'/^ m —•
OC d d d d o
taken by the grabs, there was no clear relation- +1 tl +1 +1 4.
00
m
ship between these two variables, e.g., the sam- 4 fN
fN

d 1
fM

d c d d
—1
0
pling area of the Van-Veen grab was about twice
vi
that of the Ponar grab but the volumes of subs- m c <N

trata taken by each grab were often similar or d d d d d d

+1 +1 +1 +1 +1 +1 4.
even lower for the larger Van-Veen grab (Table •0 IO r- r- WI m
1). It is also worth noting that the mean volume o c o o O o c

of substratum taken by a weighted Ponar grab OJ r- r-l ,_

was always larger than the corresponding vol- d d d d d d 0
ume for the unweighted Ponar grab, even +1 +1 +1 +1 +1 +1 +
IT, o o o fN -o
though the sampling area was obviously the d ( S -^ O o 0
same.

'.06
(N l O r- 00 00
The mean depth of penetration for each grab —— o o —
was a useful index of performance because it was 4 +1 +1 +1 +1 +1 fl +
independent of the sampling area. Mean values q q 1/1

d u-j
*" f^

were calculated directly from eaeh bite profile

(Fig. 2) by dividing the cross-sectional area by rn rN o lO

the width at the surface. They were also calcu- d d d d d d c

2-64
+1 +1 +1 +1 +1 +1 +
lated indirectly from the tank and field experi- c ta >£> q •o — l O 0 0
ca oc
ments by dividing the volume of substratum in « JZ
f^
d d d "^ f , c
each catch by the sampling area of the grab. For o
—• ca ,_,
u t
each grab, there were usually no significant dif- £
a. d d d d d d 0
00

fN
ferences between values for the profiles in sand E c +1 +1 +1 +1 +1 +1 +1
00 •o
and gravel, the tank experiments with fine gravel o u^
ectilytr

II -^ d fN d ri r i C
(2-4 mm) and the field experiments at site 1
(Table 2). Exceptions were the significantly •o
c
fN o lo u-l 0 0 --1

lower values for the Dietz-La Fond and Allan •o -- d d d d 0

• o ,-t +1 +1 +1 +1 +1 4-i +
grabs at site 1 and in the tank experiments with u U rN 0 0 q -- q 00
8-1

r^ "*' fN
-~ c
fine gravel and the significantly higher value for
±95^

u E
•c
the Birge-Ekman grab at site 1. The low values o r-l r^ l O VC r..
were due to some loss of gravel or mud from j£
2 c
M
— r-' o O o o c

both grabs and the high value for the Birge- bC u +1 +) +1 +1 +1 +1 +1

J3
E c r- fN r- - M

Ekman grab was due to its greater depth of u

ca
fN •rt "^ rN
u
penetration in mud. As substratum size M
in
(N _
increased, the mean depth usually decreased for c
U
3
o O O o o c
(N ni
C
o a
eaeh grab except for the Dietz-La Fond and +1 +1 +1 +1 >.| +1 +1
:rat

u-l r- u-l u-l c

Birge-Ekman grabs in the tank experiments. u Cu
j=

Values for the tank and field experiments show

that in mud (site 1) the mean depth of penetra- fN fN u-l

tion was >3 cm for all except the Friedinger d d O d d d d

grab and >5 cm for the Ponar and Birge-Ekman i «1
t l +1 +1 +1 +1 +1 +1
l/l CT- u-l 1—

grabs, whilst in uniform gravel (2-4 mm) the o O u-i

u
values were >3 cm for the Birge-Ekman grab X
a
and >5 cm for the Van-Veen and both Ponar CO
1 Ponar

s
-La Fond

grabs. When small stones (8-16 mm) were pres-

man

ent, none of the grabs sampled to a mean depth E c

u
inge

S w UJ
>5 cm and only the Ponar grabs exceeded a m B ca
E. J
—
^ OCI i n T3 u
mean depth of 3 cm. When larger stones (>16 ca w .£>
D
u c c U E?—
.a > 'C Q ia <
mm) were present in a predominantly gravel t/1
u.
bottom (site 2), no grabs sampled to 3 cm and

the weighted Ponar was the only grab to reach 2
cm. This inability of the grabs to penetrate even
to 3 cm on a bottom with larger stones was
confirmed by both the tank and field experi-
ments, apart from the rather odd value for the
Birge-Ekman grab on the largest substratum
size in the tank experiments.
Some notes were made on the problems of
taking samples with each grab. The Van-Veen
grab was rather heavy (21 kg) and cumbersome
with its two long arms, was difficult to set and
occasionally jammed with a stone or twig bet-
ween the jaws. Although the weighted Ponar
grab was also rather heavy (23 kg) and had to be
set carefully, it worked well and sampled consis-
tently to the greatest mean depth in most of the
tank and field experiments (Table 2). The
unweighted Ponar grab was easy to operate,
rarely failed and was light enough (14 kg) to
handle from a small boat. The Friedinger grab
was easy to operate but failed frequently when
larger stones (> 16 mm) were present and did
not function successfully at site 4. Apart from
the problem of its weight (21 kg), the Dietz-La
Fond grab jammed frequently when gravel (2-4
mm) was present. The Birge-Ekman grab was
easy to operate especially on a muddy bottom,
but jammed frequently when fine gravel (2-4
mm) was present. Frequent jamming was also
the major problem with the Allan grab on all
substrata and one small stone or twig between a
side plate and a jaw was enough to prevent clos-
ure.
Therefore, the general performance charac-
teristics of the grabs can be summarized as fol-
lows. All grabs, except the Allan grab, showed a
marked reduction in sampling area with depth of
penetration into the substratum (Fig. 2), and
usually took a representative sample of the subs-
tratum at each site (Fig. 3). The performance of
the Friedinger, Dietz-La Fond and Allan grabs
was generally poor. These grabs frequently
failed to operate and took small samples of sub-
stratum, except on a muddy bottom. The mean
depth of penetration into the substratum was
always less than 2 cm for the Friedinger grab and
less than 3 cm in all substrata except mud for the
Dietz-La Fond and Allan grabs. Although the
Van-Veen and Birge-Ekman grabs sampled to a
mean depth greater than 3 cm in both mud and
fine gravel (2-4 mm), the Van-Veen was dif-
ficult to operate and the Birge-Ekman jammed
frequently in fine gravel. Both Ponar grabs
Comparative study of grabs 107
operated well and sampled to a mean depth
equal to or greater than 5 cm in both mud and
fine gravel. They were the only grabs which
sampled to a depth >3 cm when small stones
(8-16 mm) were present and the weighted Ponar
was the only grab to attain a depth of 2 cm when
larger stones (>16 mm) were present in a pre-
dominantly gravel bottom. The weighted Ponar
always sampled to a greater depth than the
unweighted version but the latter was slightly
easier to operate. All grabs performed badly
when larger stones (>16 mm) were predomin-
ant in the substratum.
Tank experiments with plastic pellets
TTie term 'efficiency' refers to mean catch
expressed as a percentage of the numbers actu-
ally present and is synonymous with the term
'accuracy'. It must not be confused with the term
' precision" which refers to the error term
attached to the mean and is expressed as 95%
confidence limits in the present study (see Sutc-
liffe. 1979, for a more detailed discussion of
these terms). As the number of plastic pellets in
the trays was known (see methods), both mean
number m"' and efficiencies (both with 95%
CL) were estimated (Fig. 4).
Catches of pellets just below the surface of the
substratum were usually higher than the corres-
ponding catches for 'invertebrates' at a depth of
c. 3 cm, and the catches with each grab decreased
markedly as the substratum size increased (Fig.
4). In fine gravel (2-4 mm), the efficiences of
both Ponar grabs (Fig. 4b,c) were 100% for
surface pellets and about 70% for pellets at c. 3
cm. The corresponding mean values for other
grabs at the surface and a depth of c. 3 cm
respectively were, in decreasing order of effi-
ciency, 87% and 56% for the Van-Veen, 73%
and 37% for the Birge-Ekman. 51% and 36%
for the Allan, 59% and only 7% for the
Friedinger. 22% and 26% for the Dictz-La
Fond. When the substratum was small stones
(8-16 mm), the weighted Ponar was the only
grab with 100% efficiency for surface pellets
whilst the unweighted Ponar was slightly lower
at 88% and the values for all other grabs were
below 50%. The corresponding values for pel-
lets at a depth of c. 3 cm were just below 50% for
the Ponar and Van-Veen grabs, about 20% for
the Dietz-La Fond grab and less than 7% for the
Friedinger, Birge-Ekman and Allan grabs.
108 J. M. Elliott and C. M. Drake

Particle size (mm)

2-4 8-16 16-32 32-64

20000n
Surface

15000-

10000-..
jfc nlOO

50
5000-

0 A 0
c.
3cm deep
12500-1 100
10000-

50
5000-

25000-1 Total
lOO

20000-

15000-

50
10000-

5000- \

a b c d e f g o b c d e f g a b c d e f g a b c d e f g -•o

FTG. 4. Mean number per m' and percentage efficiences (with 95% CL) for plastic pellets in different substrata
with modal sizes of 2-4 mm. 8-16 mm, 16-32 mm and 32-64 mm. Separate values are given for pellets just below
the surface, at a depth of c. 3 cm and at both depths combined. Separate values are given for each grab: (a)
Van-Veen, (b) weighted Ponar, (c) Ponar, (d) Friedinger, (e) Dietz-La Fond. (0 Birge-Ekman, (g) Allan.

These decreases in efficiency were even more
marked when the substratum was slightly larger
stones (16-32 mm) and the Friedinger and Allan
grabs did not catch any pellets on this substratum
(Fig. 4d,g). Catches for all grabs were zero or
almost zero on the largest substratum (32-64
mm).
As the density of pellets in these experiments
was rather high (23142 ± 655 m"^, see
methods), it was possible that the efficiences
quoted above would be lower at lower densities.
Tliereforc, the density of pellets was reduced to
80%, 60%, 40% and 20% of their usual value of
23142 m^^ in a series of experiments with a
Ponar grab and a substratum of fine gravel. The
grab and substratum were chosen because they
provided the highest efficiencies in the original
experiments. There were clearly no significant
changes in efficiency as the density of pellets
decreased (Fig. 5). It was therefore concluded
 Comparative study of grabs 109

Surface c. 3cm Deep Totol

100-

75-

i 50-

25-

100 80 60 40 20 100 80 60 40 20 100 80 60 40 20

Density of plastic pellets (% of max values)

FIG. 5. Percentage efficiencies (with 95% CL) for a Ponar grab sampling plastic pellets at different densities
(density is expressed as a percentage of the density used in all the other tank experiments) in gravel (modaJ size 2-4
ram); separate values are given for pellets just below the surface, at a depth of t. 3 cm and at both depths combined.

that the efficiencies obtained in the more exten-
sive experiments were representative of differ-
ent population densities.
Therefore, the results obtained from Ihe exper-
iments with plastic pellets agreed with the gen-
eral performance characteristics described in the
preceding section. Efficiencies for the total
catches of the Friedinger, Dietz-La Fond and
Allan grabs were low, with values of less than
45% for fine gravel (2-4 mm), <22% for small
stones (8-16 mm), <5% for slightly larger
stones (16-32 mm) and close to 0% for larger
stones (32-64 mm). If 50% is assumed to be Ihe
minimum acceptable efficiency for total catches,
then the Ponar, Van-Veen and Birge-Ekman
grabs were adequate for a substratum of fine
gravel (2—4 mm), only the two Ponar grabs were
adequate for small stones (8-16 mm) and no
grabs were adequate in a substratum of larger
stones (>16 mm).
Field trials
Oligochaetes and larvae of chironomids were
the dominant invertebrates at each site. Most
taxa have been collated in major groups for the
analyses of this and the next section, but a full list
of taxa is given in Appendix 1 so that a
summary is available for the invertebrate com-
munity at each site. The relative abundances of
major taxa were expressed as percentages of the
total catch and were very similar for most grabs
at each site (Table 3). Major exceptions were
the Allan grab at site 2, the Dietz-La Fond grab
at sites 1 and 2 and the Friedinger grab at sites 2.
3 and 4. The mean numbers of taxa taken at site
1 were not significantly different for the Van-
Veen, Ponar, Birge-Ekman and Allan grabs, but
were significantly lower for the Friedinger and
Dietz-La Fond grabs (Fig. 6). There were two
distinct groups of grabs at sites 2 and 3 with no
significant differences between mean numbers
of taxa within each group. The Van-Veen and
both Ponar grabs were in the first group whilst
the remaining four grabs formed the second
group with significantly lower values. Mean
numbers of taxa were low but not significantly
different for all grabs at site 4, apart from the
zero value for the Friedinger grab. TTie precision
of these estimates of mean numbers of taxa was
expressed as 95% CL and was very similar at
sites 1-3 but decreased markedly at site 4 (Fig.
6).
The estimates of mean numbers of inverte-
no J. M. Elliott and C. M. Drake

ro_J " ^ ^
— rs •* —
+1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 tl tl tl tl tl tl tl tl tl +1 tl +1
(N o * r s Ov -H r - ; p I"-, - ^ t

00 -H r-^ r^ rs (S
UJ
+1 +1 +1 +1 +1 +1 +1 +1 +1 +1 t l +1 +1 +1 +1 +1 tl tl tl tl tl tl tl tl tl +1 t l tl t l
Tt H-) T t u-l r-l p ^ ON f - O>

CQ d o\ c — (*i vS
,— IJ-J r J ,—

^ fO vC ro
ro O lo 00 - - f.j

19.
^ •; r s <s -^ d

tl tl +1 tl c tl tl tl tl tl tl t[ ti tl +1 tl tl tl tl tl tl tl tl tl tl tl tl t l tl
r~ n >o I'l lo u^ ll-> ^ r- o>

5:3 r s d o6 r-
Tj- CS —I rowi

+1 +1 +1 +1 +1 +1 +1 +1 +1 +1 tl tl tl tl tl tl tl tl tl tl tl tl tt tl tl
(N o - ^ \o fo r- o o o o

ts r-^ r-i f^i ^ —^ <?v O- 1.0 —

+! +1 +1 +1 +1 +1 +1 +1 +1 tl tl tl tl tl tl +1 tl tl tl tl tl tl tl tl tl tl tl tl
<N-^rOpQOc--iiorsr-;
rsi-iro — c ' o v d o o r o
ro — _____

00 •— rs r- '-" <N
I I I I I I I I I I tl tl tt +1 tl tl tl tl tl tl tl ti tl tl tl I I I I
• » T t D O — ; - - r j - 0 O — O\

\D O^ ^ < ; <s 00 o o o\
"":^d'^'^P°'?vd"P
W Ov fO ro — ^H 00 f S •--
+1 +1 +1 +1 +1 +1 +1 +1 +1 +1 tl tl tl tl tl t l tl tl tl tl tl tl tl tl tl t l t l tl tl
00 O ^ Ov - - 'O
O\ o i d — ro 1^

« 3 o S ^ 'E .S
u -0 « 5 i«

mill II l i l l § I e
ra a p D.C £ J3 m £
ZH-JUJh-OUf-0
:2 ^
In
O
 Comparative study of grabs 111

Site I Site 2 Site 3 Site 4

60 n

40-

20- \h

0'
70000-

6000OH

40000-

20000-

3-T

^"

1- r-

Q c d e f g a b c d e f g a b c d e f g a c d e f g

FIG. 6. Mean numbers of taxa per sample, mean numbers of invertebrates per m'and mean indices of diversity (all
with 95% CL) for different grabs at sites 1-4: (a) Van-Veen, (b) weighted Ponar, (c) Ponar, (d) Friedinger, (e)
Dietz-La Fond, (f) Birge-Ekman, (g) Allan.
112 J. M. Elliott and C. M. Drake
brates m ^ varied considerably between grabs at
all sites and the precision of these estimates was
often poor, especially at site 4 (cf. 95% CL in
Fig. 6). There were no significant differences
between values forthe Ponar, Birge-Ekman and
Allan grabs at site 1, but values for the Van-
Veen, Friedinger and Dietz-La Fond grabs were
significantly different and values for the
the weighted Ponar grab was significantly higher
than the mean values for all the other grabs,
values for the Van-Veen, unweighted Ponar,
Dietz-La Fond and Birge-Ekman grabs were not
significantly different, and values for the
Friedinger and Allan grabs were significantly
lower than all other values. There were no signif-
icant differences between values for nearly all
grabs at sites 3 and 4, the exceptions being the
significantly lower values for the Friedinger and
Allan grabs at site 3 and obviously the zero value
for the Friedinger grab at site 4.
An index of diversity was also calculated for
each sample and the non-parametric informa-
tion statistic (//) of Shannon & Weaver (1949)
was used. Although this index is not without its
critics (e.g., Taylor. Kempton & Woiwod,
1976), it has been claimed by others to be a
suitable index for comparing the diversity of
invertebrate communities in rivers (see refer-
ences in Weber, 1973; Hellawell, 1978). The
indices of diversity followed a pattern similar to
that shown by the number of taxa (Fig. 6). but
the indices were much more similar than the
corresponding values for taxa, especially at site
2. Therefore, the indices of diversity were rather
disappointing because they did not add any
information to the conclusions already made
from the comparisons between mean numbers of
taxa and numbers m^'.
Therefore, the results obtained with live
invertebrates from the field trials generally
agreed with those obtained from the tank exper-
iments with plastic pellets and with the general
performance characteristics, especially the ina-
bility of some grabs to penetrate very deeply into
the substratum (cf. values in Table 2 and Fig. 6).
The relative abundances of major taxa were very
similar for most grabs at each site, but the
Friedinger and Dietz-La Fond grabs were once
again the major exceptions. In terms of both
mean number of taxa and mean number of
invertebrates per m^ the performances of the
Ponar, Birge-Ekman and Allan grabs were high
on the predominantly muddy bottom at site 1.
The Van-Veen grab provided a comparable
estimate for the number of taxa but a disappoint-
ingly low value for number m' \ whilst the per-
formances of the Friedinger and Dietz-La Fond
grabs were poor at site 1. The Van-Veen and
Ponar grabs provided similar estimates of mean
number of taxa on the predominantly gravel bot-
tom with some larger stones (> 16 mm) at site 2,
but only the weighted Ponar grab provided an
estimate of mean number m"^ that was signific-
antly higher than the values for the other grabs.
It was therefore concluded that the weighted
Ponar was the only grab to approach an adequ-
ate pjerformance at this site. The performances
of the Friedinger and Allan grabs were excep-
tionally poor at sites 3 and 4, especially for esti-
mates of mean number m"^ As the perfor-
mances of the other grabs at sites 3 and 4 were
rather similar with very poor precision (cf. the
confidence limits in Fig. 6) and it has already
been shown that the mean depth of penetration
into the substratum was less than 0.8 cm (Table
2), it was concluded that all grabs performed
badly when larger stones (>16 mm) were pre-
dominant on the bottom.
RelatioiLship between the mean and variance, and
the coefficient of variation
The relationship between the arithmetic mean
{^) and the variance {s^) of samples of inverte-
brates frequently follows a power law (Taylor.
1961; Taylor, Woiwod & Perry. 1978):
(1)
where a and b are constants. In the tank experi-
ments, eqn 1 was found to be an excellent des-
cription of the relationship between the var-
iances and means for the catches with the differ-
ent grabs. Values of the constants a and b were
not significantly different for the catches of plas-
tic pellets just below the surface, at a depth off.
3 cm and at both depths combined (Table 4. Fig.
7a). In the field trials, values of a and b were
calculated for major taxa and total numbers at
each site as well as for total numbers at all sites.
Equation 1 was an excellent fit for all except two
taxa at site 4 (Table 4) and the reiationship for
total numbers at all sites is illustrated in Fig. 7b.
Significant values of the power b in eqn 1 varied
from 1.14 to 2.34 and this range is well within
the range obtained for terrestrial invertebrates
 Comparative study of grabs 113

TABLE 4. Values of the constants a andfc ± 95% CL in eqn 1 for various taxa at
sites 1-4 and for tank experiments with pellets (n = number of values for each
regression analysis; r' = coefficient of determination; significance levels indicated
by asterisks:* F<0.05; " P<O.OI; NS = not significant, /*>0.05)

h ± 95% CL

Pitidium 6 3.268 I.fi79 ± 0.301 0.984

Naididae 6 5.402 1.694 ±0.536 0.951
TubiHcidae b 1.789 1.883 + 0 943 0.885
Copcpoda 6 2.068 t.874 ±0,152 0.997
Asetlus aquaiicia 6 2.822 1.770 ±0.591 0.945
Sialis lularia 6 -0.194 1.814 ±0.581 0.949
Tanypodinae 6 2.138 1.589 ±0.334 0.978
Oiironomini 6 0.827 1.932 ±0.255 0.991
Ta n y tarsi ni 6 2.171 1.803 ±0.307 0.985
Tdlal numtiers 6 2.239 1.75l>± 0.381 0.976

Naldidaf 7 2.278 1.823 ±0,32.1 0.977

Lumbriculidae 7 2.192 1,585 ± LOOl 0.768
Tubificidae 7 31,542 1.391 ±0.558 0.892
Ephemrrella igniia 7 -0.167 1,825 ± 0.258 (1.983
Anabolia nerva^a 7 1.941 IJ68 ±0,412 t).M6
Tanypodiiiae 7 1.435 r6S0 ± 0,476 0.M3
Diamesinue 7 2.390 1.137 ± 0.407 0.912
Orthoctadiinae 7 0.S46 2.037 ± 0.523 0.952
Chi rono mini 7 2.813 1,703 ± 0,440 (1.952
Tanytarsini 7 0,936 1.921 ± 0.530 0.946
Total numbers 7 17.149 1.493 ±0.519 0.916

Naididae 7 0,602 2.166 ±0.436 0.970

Tubjficidai: 7 1.435 3.0.19 ± 0.136 0.997
Ephtmirelta ignilti 7 2.756 1.239 ±0,628 0.837
Anabolia nervosa 7 I 996 1.516 ± 0,401 0,950
Orthocladiinae 7 2,015 1.754 ± 0.363 0.969
Chironomini 7 0.946 2.104 ± 0,218 0,992
Tanylamni 7 1.85(1 1.844 ± 0,382 0.969
Tiilal numbers 7 0.442 2,119 ± 0 3 9 3 0.975

Naididae 5 0 229 2.342 ± 0.856 0.96Z

Tanypodinae 5 1,003 2.273 ±2.156 0.790
Corynoneurini 5 0.803 2.156 ±4.027 0.492
Diamcsinae 5 2,970 1.609 ±0.932 0,910
Orthocladiinae 5 3.597 1.798 ±0,310 0.991
Chironomini 5 0.506 2.553 ± 2,732 0.747
Tanytarsini 5 1,813 1.937 ± 1.351 0.874
5 0.428 2.134 ±0.722 0.967
Total numbers

Alt sites
25 1.864 ±0.207 0,938
ToUl namb«n

Tank experiments
24 3.375 1.509 ±0.110 0.973
Surface 19 3.715 1.446 ±0.017 0,951
c, 3 cm deep 24 3311 1.516 ± 0.109 0.974
Talal
114 J. M. Elliott and C. M. Drake

500000 r (g ,

100000

2000000 (b)
10000
[000000

1000

100000
100

10000

1000
•I if
0-1 I 1 I I I 500 J 1 I I IILlJ
0-1 100 1000 2000 10 100 1000 2000

FIG. 7. Log-log plot of the relationship between the variance (i') and the mean {x) for all grab samples: (a) on
different substrata in the lank experiments with values for pellets just below the surface (•), at a depth off. 3 cm
(x)andat botb depths combined (O);(b) total numbersof invertebrates in the field samples with values for site 1
(#), site 2 (O), site 3 (+) and site 4 (x). Regression lines are fitted to the data.

(see appendix tables in Taylor, Woiwod & Perry.
1978). Therefore, the variahihty in each sample,
and hence the precision of the sample mean, was
chiefly related to the mean catch rather than the
type of grab used at each site.
The coefficient of variation (CV) is often used
to compare the precision of samplers and is sim-
ply the standard deviation {s) expressed as a
percentage of the sample mean {CV = 100s/.S).
If eqn 1 is a suitable model for the relationship
between s^ and .r, then the relationship between
CV and jc is simply derived from eqn 1 as (see
Appendix 2):
and increase for b>2. In both cases. CV is
related to .r andisindepenent of Jc only whenfc=
2. The effect of variations in the sample mean on
the value of CV will become more important as
the difference between the value of b and 2
increases. It is obvious from the values of b
obtained in the present study (Table 4) that this
effect will be highly significant for many taxa and
three examples for total catches are given in Fig.
8. It is therefore concluded that the coefficient of
variation is an unreliable statistic that cannot be
used to compare the precision of grabs.

CV = (2) Discussion

where a and b are constants with the same values Information on the performance of marine sam-
as the constants in eqn 1. It can be seen from eqn plers has been summarized by Holme & Mcln-
2 that as x increases, CV will decrease for b<2 tyre (1971). Heavy marine grabs, such as the
 Comparative study of grabs 115

TABLE 5. The tiutnber («) of sampling units per sample required for different levels of
precision (95 % CL expressed as a percentage of the sample mean), for different values of the
sample arithmetic mean (x) (N.B.i^ is the mean number per sample, not mean number m ^) and
for different values offcin eqn 4; the value of/was 2 for n >25 but more exact values were used
forn <25. The value of a in eqn 4 is assumed lobe one for this table but \{a ^ I, then multiply
the appropriate value of f? by a to obtain the correct estimate of n

Mean number per sample (x)

(%) 0.5 I 5 10 20 50 100 1000

b= 1
10 800 400 80 40 22 10 6 1
95% CL 20 200 100 22 12 7 3 1 1
as 40 50 26 7 4 1 I 1 I
%ofi 60 24 13 3 1 1 I 1 1
80 14 8 1 1 I 1 1 I
100 10 6 1 1 1 1 1 1

b - L5
10 566 400 178 126 90 56 40 15
95% CL 20 140 100 45 32 24 16 12 5
as 40 35 26 13 10 8 6 5 1
% of.; 60 17 13 7 5 4 2 1 1
80 11 8 5 3 2 1 1 1
100 8 6 2 2 1 1 1 1

b = 2.0
10 400 400 400 400 400 400 400 400
95% CL 20 100 100 100 100 100 100 100 100
as '40 26 26 26 26 26 26 26 26
%oix 60 13 13 13 13 13 13 13 13
80 8 8 8 8 8 8 8 8
100 6 6 6 6 6 6 6 6

b = 2.5
10 283 400 894 1265 1789 2828 4000 12649
95% CL 20 70 100 224 316 447 707 1000 3162
as 40 19 26 56 79 112 177 250 791
% of? 60 10 13 26 35 50 79 111 351
80 6 8 14 20 28 44 63 198
100 4 6 9 13 18 28 40 126

Van-Veen weighing 45-60 kg and the Smith-
Mclntyre weighing 45 kg, will penetrate into the
substratum to a mean depth of ^8.5 cm in sand
and ^13 cm in soft mud but values of 3-8 cm are
most frequent. The latter values are therefore
comparable to those recorded for some of the
grabs sampling in mud and fine gravel in the
present study (Table 2). Few comparisions can
be made with previous comparative work on
samplers for freshwater benthos because all the
following investigations are for lake benthos and
only two studies have compared more than three
samplers.
Although he made no direct comparisons
between Ponar and Birge-Ekman grabs, Hud-
son (1970) compared both grabs with an
orange-peel grab in a lake with a bottom of field
soil covered by variable levels of silt' and conc-
luded that the Ponar grab provided the highest
estimates of numbers m"^^ Howmiller (1971)
also found that the Ponar grab performed better
than the Birge-Ekman grab on hard, muddy
sand in a bay of a lake, but the Birge-Ekman
grab was superior in softer, muddy sand prob-
ably because it created a smaller shock wave and
penetrated deeper into the softer sediments. Sly
(1969) summarized in general terms the results
of field trials in Lake Ontario with three gravity
corers, two multiple corers and the Birge-
Ekman, Ponar, Shipek, Dietz-La Fond,
Franklin-Anderson and Petersen grabs.
Although few results were given, the general
116 J. M. Elliott and C. M. Drake

300

100

50

M L 111!
0-1 0-5 10 50 100 500 1000 2000

200 (c

100

J I I I I
10 30 100 500 1000 2000

J I I I I 1 I I I

too 500 1000 1500

FIG. 8. Log-log plot ofthe relationship between the coefficient of variation {CV) and ihe mean {x) for: (a) ali grabs
sampling on different substrata in the tank experiments with values for pellets just below the surface (•),ai depth
of c. 3 cm (X) and at both depths combined (O);(b) Ponar grab sampling pellets at different densities (symbols arc
the same as in (a)]; (c) total numbers of invertebrates for all yrabs in the field samples with values for site I (•), site
2 (O), site 3 (-(-) imd site 4 (x). Regression lines arc fitted to the data.

conclusions were the same as those in a more
detailed report by Flannagan (1970) who com-
pared the sampling abilities of two corers with
various modifications, two multiple corers and
six grabs in Lakes Ontario and Winnipeg. The
grabs were the same as those used in the previ-
ous study except that a tall Birge-Ekman grab
was used instead of a Petersen grab. Smith-
Mclntyre, orange-peel and Petersen grabs were
not included in these trials because it had already
been shown that the Smith-Mclntyre grab was
superior to the orange-peel and Petersen grabs
(Beeton. Carr & Hiltunen. 1965) and that the
Ponar grab was at least as efficient as the
Smith-Mclntyre grab and was less subject to
mechanical failure (Powers & Robertson,
1967). The Ponar grab and Shipek scoop were
the only samplers to function on a gravel bottom
and they, together with the Franklin-Anderson
grab, were the only samplers to function on a
sandy bottom. Estimates of population densities
m"^ were similar for the Shipek and Ponar samp-
lers on both sand and gravel, but were signific-
antly higher for the Franklin-Anderson grab on
sand. The performances of the Shipek and
Franklin-Anderson grabs were poor on a muddy
bottom when compared with the Ponar grab,
both Birge-Ekman grabs and one of the multiple
corers. A standard Birge-Ekman grab and a
multiple corer were the only samplers to provide
population density estimates that were similar to
those obtained from cores taken in the mud by a
diver. The best general purpose samplers were
the Ponar grab and Shipek scoop, but the per-

formance of the latter was poor in mud and it was
rather heavy (70 kg) for operation from a small
boat.
The general conclusions from these studies
agree with those of the present study. Ponar,
Birge-Ekman and Allan grabs performed well
on a predominantly muddy bottom (particle size
0.004-0.06 mm), their estimates of mean num-
bers m"^ were significantly higher than those
obtained from the other grabs and the mean
depth of penetration into the mud exceeded 5
cm for the Birge-Ekman and Ponar grabs. When
the substratum was fine gravel (modal size 2-4
mm), efficiencies in terms of numbers m'^
exceeded 50% for the Van-Veen (71 %), weigh-
ted Fonar (93%), unweighted Ponar (94%) and
Birge-Ekman (54%) grabs and the mean depth
of penetration exceeded 5 cm for all these grabs
except the Birge-Ekman. A weighted Ponar
grab was the only sampler to perform adequately
on a predominantly gravel bottom with some
larger stones (>16 mm) present. The general
performance of the Friedinger and Dietz-La
Eond grabs was poor. When larger stones (>16
mm) were predominant on the bottom, all grabs
performed badly with efficiencies below 12% in
the tank experiments and a mean depth of
penetration into the substratum of less than 0.8
cm in the field experiments.
It has already been shown that the coefficient
of variation (CV) is an unreliable statistic for
comparing the precision of grabs. Unfortu-
nately, other workers have used it for this pur-
pose, e.g., Weber (1973) used CV values to
compare the performances of Birge-Ekman,
Petersen and Ponar grabs on mud, sand and
gravel. If eqn 1 is a suitable model for the rela-
tionship between the variance and mean for a
series of samples, then the CV can be used to
compare the precision of grabs only when the
mean catch is the same for each grab or when the
value of b in eqn 1 is equal to two.
Equation 1 can also be used to estimate the
number (n) of replicate sampling units required
for different levels of precision. If the 95% CL
are expressed as a percentage of the arithmetic
mean (Jc), then the index of precision (D) is
simply this value divided by 100. For example, if
we can accept 95% CL of ±40% of the mean
(equivalent to a standard error of about 20% of
the mean), then D = 0.4. The sample size (n
sampling units) needed to obtain an estimate of
the population mean with a known level of preci-
Comparative study of grabs 117
sion is given by the following equation (from
Elliott, 1977):
D'i' (3)
where / is found in tables of Student's
/-distribution, D is the index of precision,.v' is the
sample variance and f is the sample mean. From
eqns I and 3, we obtain the following equation
(see Appendix 2):
= I- ax
Tt> -2 n~i
(4)
where a and b are constants with the same values
as the constants in eqn 1. Equation 4 has been
used to calculate n for various combinations of J,
b and D (Table 5). The range of values for x is
0.5-1000 and this range probably covers most
values obtained in field samples (note that x is
the mean number per sample, not an estimate of
mean number m"^). Most values of b lie in the
range 1.0-2.5 (e.g.. values in Table 4) whilst the
required values of D will usually lie well within
the range O.l-I.O (10-100%). fhe estimates of
n in Table 5 are for a = 1. If this assumption is
incorrect, then the value of n in the table must be
multiplied by the value of a to obtain the correct
estimate of n.
It is obvious from Table 5 that for the same
value of JC, the value oin decreases as the preci-
sion decreases, i.e.. D increases. When i = I,
values of « are similar for all values of 6. AsJc
increases for a constant value of £), the value oin
decreases \ib<2, increases if 6>2 and remains
constant for b = 2. Therefore, as the spatial
distribution of the invertebrates becomes more
clumped and b increases, the relationship bet-
ween n and mean density does not remain cons-
tant. Reasonable levels of precision (95% CLof
20-40% of the mean) are possible with rela-
tively small samples (n<30) whenfc<2 andi>5,
but these sample sizes may have to be doubled or
trebled when o is 2 or 3, respectively, and such
values offlare probably common (e.g., values in
Table 4). More information is clearly needed on
the values of o and b and their ranges for various
freshwater taxa, for different life stages of the
same species and for different types of habitat.
When this information is available, it will be
possible to predict with greater accuracy the
sample sizes required for different levels of pre-
cision and different taxa.
118 J. M. Elliott and C. M. Drake
Acknowledgments
We wish to thank Mrs P. A. Tullett, J. D.
Allonby, A. P. Martin and F. R. Ohnstad. for
their assistance with ihis work. I. R. H. Allan, for
the loan of his grab, and Dr D. W. Sutcliffe, for
his constructive comments on the manuscript.
Some of the cost of the research has been met by
the Department of the Environment (Contract
No. DGR 480/329).
References
Allan I.R.H. (1952) A hand-operated quantitative
grab for sampling river beds. Journal of Animal
Ecology, 21, 159-160.
Beeton A.M., Carr J.F. & Hiltunen J.K. (1965) Sampl-
ing efficiencies of three kinds of dredges in Southern
Lake Michigan. Proceedings of the 8th Conference Lisitsyn A.P. & Udintsev G.B. (1955) New model
on Great Lakes Research, 209. Iniernational
Association for Great Lakes Research. Michigan.
Bhaud M & Duchfine JjC. (1977) Observations sur
Icfficacitt comparde tie deux benncs. Vie et Milieu
iSerieAl 27. 35-53.
Birge E.A. (1921) A second report on Hmnological
apparatus. Transactions ofthe Wisconsin Academy
of Sciences, Arts and Letters, 20, 533-552.
Birkett L. (1958) A basis for comparing grabs, yourfia/
du Conseil, 23, 202-207.
Ekman S. (1911) Neue Apparate zur qualitativen und
quantitativen Erforschung der Bodenfauna der
Seen. Inlernalionale Revue der gcmmten Hyd-
rohiologie und Hydrographie, 3, 553-561.
Elliott J.M. (1977) Some methods for the statistical
analysis of samples of benthic invertebrates. Scien- Reineck H.E. (1963) Der Kastengreifer. Natur und
tific publications of the Freshwater Biological
Association No. 25, 2nd edition, 156 pp.
Elliott J.M., Drake CM. & Tullett P.A. (1980) The
choice of a suitable sampler for benthic macro-
invertebrates in deep rivers. Pollution Report ofthe
Department ofthe Environment U.K., (in press).
Elliott J.M. & Tullett P.A. (1978) A bibliography of
samplers for henthic invertebrates. Occasional Publi-
cations of the Freshwater Biological Association
No. 4, 61 pp.
Flannagan J.F. (1970) Efficiencies of various grabs
and corers in sampling freshwater benthos, yourna/
of the Fisheries Research Board of Canada 27,
1691-1700.
Gallardo V.A. (1965) Observations on the biting pro-
profiles of three bottom samplers. Ophelia, 2,
319-322.
Hartman O. (1955) Quantitative survey of the ben-
thos of San Pedro Basin, Southern California. Part
1. Preliminary results. Allan Hancock Pacific
Expedition 19, 185 pp.
Hellawell J.M. (1978) A comparative review of
methods of data analysis in biological surveillance. Taylor L.R., Woiwod LP. & Perry J.N. (1978) The
Pollution Report ofthe Department ofthe Environ-
ment U.K.. No. 3,45-57.
Holme N.A. (1949) A new bottom sampler,./oMrna/o/
the Marine Biological Association ofthe U.K., 28,
323-332.
Holme N.A. (1953) The biomass of the bottom fauna
in the English Channel off Plymouth. Journal ofthe
Marine Biological Association of the U.K.. 32,1-49.
Holme N.A. & Mclntyre A.D. (1971) Mef/io(i^/or r/itr
.study of Marine Benthos. IBP Handbook No. 16.
Blackwell. Oxford. 334 pp.
Howmiller R.P. (1971) A comparison of the effective-
ness of Ekman and Ponar grabs. Transactions ofthe
American Fisheries Society, 100, 560-564.
Hudson P.L. (1970) Quantitative sampling with three
benthic dredges. Transactions of the American
Fisheries Society. 99, 603-607.
Hydro Products (1973) The Shipek sediment sampler.
Hydro Products Special Bulletin, 3 pp.
Kutty M.K. & Desai B.N. (1968) A comparison of the
efficiency of the bottom samplers used in benthic
studies off Cochin. Marine Biology. 1, 168-171,
La Fond EC. & Dietz, R.S. (1948) New snapper-type
sea floor sediment sampler. Journal of Sedimentary
Petrology. 18, 34-37.
dredges. (In Russian.) Trudy Vsesoyuznogo gid'
robiologicheskogo obshchestva. 6, 217-222.
Murray T.D. & Charles W.N. (1975) A pneumatic
grab for obtaining large, undisturbed mud samples:
its construction and some applications for measur-
ing the growth of larvae and emergence of adult
Chironomidae. Freshwater Biology, 5, 205-210.
Naumann E. (1925) See und Teich. Handbuch
BiologLichen Arbeitsmeihoden, 9, 103-138.
Pettijohn F.J. (1957) Sedimentary rocks. 2nd ed.
Harper & Brothers. New York. 718 pp.
Powers C.F. & Robertson A. (1967) Design and
evaluation of an all-purpose benthos sampler. Spe-
cial Report ofthe Great Lakes Research Division,
University of Michigan, 30, 126-131.
Museum. 93, 102-108.
Reys J.P. (1964) Les priiltvemnets quantitatifs du
benthos de substrats meubles. Terre Vie. 94-105.
Shannon CE. & Weaver W. (1949) The mathematical
theory of communication. University of Illinois
Press, Urbana.
Sly P.G. (1969) Bottom sediment sampling. Procwd-
ings of the !2th Conference on Great Lakes
Research. 883-898. International Association for
Greai Lakes Research, Michigan.
Smith W. & Mclntyre A.D. (1954) A spring-loaded
bottom sampler. Journal of the Marine Biological
Association ofthe U.K.. 33, 257-264.
Sutcliffe D.W. (1979) Some notes to authors on the
presentation of accurate and precise measurements
in quantitative studies. Freshwater Biology. 9,
397-402.
Taylor L.R. (1961) Aggregation, variance and the
mean. Nature, 189, 732-735.
Taylor L.R., Kempton, RA. & Woiwod I.P. (1976)
Diversity statistics and the log-series model.ioMrna/
of Animal Ecology. 45, 255-272.
density dependence of spatial behaviour and the
rarity of randomness. Journal of Animal Ecology,
An, 383-406.

Thamdrup H.M. (1938) Der van Veen-Bodengreifer.
Vergleichsversuche iJber die Leistungsfahigkeit des
van Veen und des Petersen Bodengreifers. Journal
du Conseil. 13.206-212.
Thorson G. {1957) Sampling the benthos. Memoirs of
the Geological Society of America, 67, 61-73.
Ursin E. (1954) Efficiency of marine bottom samplers
of the van Veen and Petersen types. Meddelelser fra
Danmarks Fiskeri- og Havunders0gelser, 1, No. 7,8
pp.
Van Veen. J. (1936) Onderzoekingen in de Hoofden.
in verband met de gesieldheid der Nederlandsche
Kust. Dissertation, University of Leiden.
Weber C.I. (1973) Biological field and laboratory
methods for measuring the quality of surface waters
and effluents. Environmental Monitoring Series,
U.S. Environmental Protection Agency, EPA
670/4.73.001.
Welch P.S. (1948) Limnological Methods. McGraw-
Hill Book Co. Inc.. U.S.A. 381 pp.
Wigley R.L. (1967) Comparative efficiencies of van
Veen and Smith-Mclntyre grab samplers as
revealed by motion pictures. Ecology. 48,168-169.
(Manuscript accepted 19 February 1980)
Appendix 1
Taxii recorded at the four sites ( 1 . 2. 3 and 4): taxa in
square brackets were grouped together in the analyses
described in the text.
MoUusca: Ancylus fluviatilis Mull. 1-4; Lymnaea
peregra (Mlill.) 1. 2; Planorbis ulbus MQll. 1. 2; P.
contortus (Linn.) 1. 2:Potomopyrgu.s jenkinsi (Smith)
l~A;Segmenlina complanaia (Linn.) 1; Valvata crislata
Miill. 1; V. macrostoma Steen. 1; Bivalvia 1, 2, 4;
[Fisidium casertanum <PoIi) 1, 2, 4; Sphaerium cor-
neum (Linn.) 1, 2; S. lacustre (Mull.) 1].
Hirndinea: Erpobdella octoculata (Linn.) 1-4; Glos-
siphonia comptanaia (Linn.) \-A; Helobdellastagnalis
(Linn.) 1-4; Hemictepsis marginata (Mull.) 1; Pis-
cicola geometra (Linn.) 3; Theromyzon tessulatum
(MQU.) 1.
OUgochaeta: Tubificidae 1-3 [Aulodrilus pluriseta
(Piguet) t. 3; Bothrioneurum vejdovskyanum Stole 2;
Limnodrilus hoffmeisteri Claparfede t - 3 ; Peloscolex
ferox (Eisen) 1, i; Poiamoihrix Vejd. & Mrazeksp. 1;
Rhvacodrilus coccineus (Vejd.) 3; Tubifex tubifex
(MUn.) 1, 3] Naididae \-A [Chaetogaster von Baer sp.
1, 3, 4; Nais barbata (Miill.) 2-^\ N. bretscheri
Michaelson 3; N. pardalis Piguet 3; N. pseudobtusa
Piguet ?>. OphidonaLt sepentina (Mlill.) \. Pristina km-
gisela Ehrcnberg 2. 3; Slavina appendiculata
(Udekem) \.2:Stylaria lacuslris {Unn.) XA.Vejdovs-
kyetla comata (Vejd.) 1. 4 | Lumbriculidae 1-4 \Lum-
briculus- variegatus (Mull.) 1-4; Stylodrilus hering-
ianus Claparfede 2, 3] Enchytraeidae 2.
Other invertebrates (except insects): Hydra Linn. sp.
1-4; Dendrocoelum lacteum (Mull.) 4; Polyeelis t - 4 ;
Comparative study of grabs 119
[P. felina (Dafyell) 3; P. lenuis Ijima I-4J Nematoda
1-4; Cladocera 1—1 [Daphnia hyalina Leydig 1;
Eurycercus lamellatus (Mlill.) 1, 2, 4; Simocephalus
vetulus (Mull.) I]; Leptodora kindli (Focke) 1;
Copepoda 1-4 [Cyclops albidus (Jurinc) l|; Harpac-
ticoida 1-3; Oslracoda 1-4 [Candona Baird sp. 1;
Cyridopsis Brady sp. 3; Herpetocypris reptans (Baird)
I: Notodromas monacha (Mull.) l\, Asellus aquaticus
(Linn.) t - 4 ; Gammarus pulex (Linn.) 1, 2, 3; Hyd-
racarina 1-4; Tardigrada 2.
insecta
Epfaemeroptera: Baetis rhodani (Pict.) 1, 3, 4; B.
scambus Etn. 4; Caenis rivulorum Etn. 2-4; Cenlrop-
tilum luteolum (Mull.) 3; Cloeon dipierum (Linn.) 1;
Ecdyonurus Eaton sp. 1; Ephemera danica Mull. 2;
Ephemerella ignita (Poda) 2-4; Procloeon bifidum
Bengtss. 2, 3.
Plecoptera: Leuctra genicuiata (Stephans) 2. 3; L.
hii>popus (Kempny) 2, 3; L. inemus Kempny 3. i
Hemiptera: Callicorixa ?praeusta (Fieb.) 1; Sigara dis-
/i>H:M(Fieb.) \;S.dorsatis L&ach l;S.falleni{Fieb.) 1;
nymphs indet I.
Coieopteni: Deronectes? elegans Panz. 1; D. sanmarki
Sahib. 2; Elmis aenea (Mull.) 1, 3, 4; Esolus paral-
lelepipedes (Mlill.) 2. 3; Gyrinus Geoffroy sp. 1-4;
Haliplus Latreille spp. 1-4; Haliplus fulvus Fabr. 1;
Hyphydrus ovaius Linn. 1; Limnlus volkmari (Panz.)
1-3; Oulimnius ruberculatus (Mlill.) 2-4; Plaiambus
maculatus Linn. 3, 4; Rhantus Lacordaire sp. 1.
Tricfaoptera: Agapetus deticatulus McLachlan 3;
Anabolia nervosa (Curtis) 2-^, Athripsodes aterrimus
(Sleph.) 2, 3; A. cinereus (Curtis) 2, 3; Dritsus
annulaius Stephans 3; Glossosoma boltoni (Curtis) 2,
3\Goerapitosa (Fabr). 2, 3: Hydropsyche Pictclsp. 3,
4; Hydroptila Dalmann sp. 4; Lepidosioma hirtum
(Fabr.) 2, 3; Motanna anguxtaia Curtis 2; Mystacldes
azurea (Linn.) 2. 3; M. nigra (Linn.) 2. 3; Plectroi-
nemia? genicuiata McLachlan 1; Polycenrropus
flavomaculatas Pictet 1, 3.4; Poiamophylax cingulatus
(Stephans) 2, 3; Rhyacophila dursaiis (Curtis) 2. 4;
Sericostomapersonatum (Spence) 1-3: Tinodes waen-
eri (Linn.) 2-4.
Diptera, Cbironomidae: Tanypodinae 1-4 [Ablabes-
myia Johannson sp. 1-3; Macropelopia Thien. sp. 2, 3;
? Natarsia Fittkau sp. 2; Procladius choreus (Meigen)
gp. sp. 1-3; Thienemannimyia Fittkau gp. spp. 2, 3;
Zavrelimyia Fittkau gp. sp. 1]; Diamesinae 2-4 [Pot-
thastia gaedii (Meigen) 2-4; P. longimana Kief. 2-4];
Orthocladiinae 1-4 [Brillia modesta (Meigen 1, 3;
Cricotopus?albiforceps (Kief.) 4; C. bicinctus
(Meigen), 2, 4; C. festiveltus (Kief.) gp. 2-A\ C. sylves-
His (Fabr.) gp. 1; C. tremulus (Linn.) gp. 2-4; C.
triannulalus (Macquart) 4;Cricotopus spp. indet. 2-4;
Corynoneura Winnertz sp. 2-4; Eukiefferiella ?bre-
vicalcar (Kief.) 2, 4; £. claripennis (Lundbeck) 4; E.
clypeata (Kief) 2-4; E. clavescens (Edwards) 2; E.
devonica (Edwards) 4; £. discoloripes Goetghebuer 4;
] 20 J. M. Elliott and C. M. Drake

E. iUdeyensis (Edwards) 2. 4; Heierotanytarsiis

apicatis (Kief.) I; Heteroirisso cladius (Sparck) sp.
1-4; Microcricotopus rectinervis (Kief.) 3. 4; Orthoc- then s = VaP~= a"' ^ " (lb)
ladius Van der Wulp s.str.spp. 2, 4; Prodiamesa
olivacea (Meigen) 2-4; Psectrocladius limbaiellus wheres ts the sample standard deviation. The relation-
(Holmgren) 1; Ps. obvius (Walker) 1; Rheocricotopus ship between the coefficient of variation (CV) and the
Thienemann sp. 1, 2, 4; Synorthocladius semivirens sample mean {x) is therefore derived as follows:
(Kief.) 1-4; Thienemanniella Kief. sp. 2—4] Chirono-
mini 1-4 [Chironomus Meigen spp. 1-4; Demicryp-
tochironomus vulneratus (Zetterstedt) 3; Einfeldia CV = \QQslx =
pagana (Meigen) 1; Endochironomus Kief. I; Glyp-
totendipes Kief. spp. 1-4; Limnochironomus nolatus and by substituting eqn lb fors, eqn 2 is obtained:
(Meigen) 1, 4; Microtendipes Kief. sp. 1-4; Paraten-
dipes Kief. sp. 1, 2; Phaenopsectraflavipes(Meigen) 2;
Polypedilum Kief. spp. 1-4; Sergentia Kief. sp. 2-4;
Paracladopelma Hamiseh sp. 2, 3; Stictochironomus CV =
Tanytarsini 1—4 \CludoHtnyiarsus manciis (Walker)] (2)
1-3; C. ni^rovittatus Goelghiiehuer 1; Micropst-ctra
Kief. spp. 1-4; Paratanytarsus Bause spp. 1-4; The relationship between the sample size {n sampling
Rheotanvlarsus pentapoda Kief. 4; Tanytarsus van der units) and the index of precision (D = 95% CL expres-
Wulp. spp. 1^1 sed as a percentage of the arithmetic mean x) is given
by:
Other Diptera: Tipulinae sp. 4; Helius Saint-Fargeau
& Serville sp. 1; Hemerodroma Meigen sp. 2;
Brachycera sp. 1; Ephydridae 1; Empidae spp. 2, 4;
Simulium Lalreillesp.4;5.flurewmFries4;5. ornolum (3)
Meigen 4; S. sublaciWre Davies 4; Ceratopogonidae
sp. l~3[Bezzia Kief. sp. l-3;C«//ct>(i/e5 Latreille sp. 3;
Dasyhelia Kief. sp. 1] Psychodidae sp. indet. 1, 3, 4
[Pericoma Walker sp. 1] where t is found in tables of Student's (-distribution.
Equation 4 is obtained by substituting eqn la for5' in
eqn 3 thus:
Other insects: Sialis luiaria (Linn.) 1,4; Coenagriidae
1.
n = t'ax'^x"^ D-^
Appendix 2
= t'ax'""'' D-' (4)
Derivations of eqns 2 iind 4.

If the relationship between the arithmetic mean (x) where a and h are constants with the same values as the
and the variance (s^) of a series of samples is given by: constants in eqn la.