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Population structure and growth of

Donax trunculus (Bivalvia:
Donacidae) in the western
Mediterranean
Maria Ramon
Marine Biology

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Marine Biology (1995) 121: 665-671
M. Ram6n 9R Abelld 9C. A. Richardson
Population structure and growth of Donax trunculus (Bivalvia: Donacidae)
in the western Mediterranean
Received: 18 August 1994 / Accepted: 19 October 1994
Abstract A comparison of shell growth in Donax trun-
culus (collected between 1988 and 1990 of Cullera, Spain)
has been carried out using an analysis of cohort progres-
sion in monthly length frequency distributions, hyaline sur-
face shell growth rings and internal microgrowth bands. In
the Mediterranean there are two periods of recruitment of
D. trunculus, one in the summer (July to September) and
the other in winter (December to February). Clams re-
cruited to the population in winter display a clear cessa-
tion in shell growth during the following summer which
may possibly be correlated with spawning, whereas indi-
viduals of the summer recruited cohort show no growth
cessation the following summer and continue to deposit
shell during this period. The normally opaque shell of
D. trunculus reveals the presence of translucent hyaline
growth rings when the shells are backlit by a strong light
source, and these have been shown to be laid down in the
shell during summer months. Formation of a hyaline ring
is accompanied by a narrowing of the microgrowth pat-
terns present in shell sections. Both the hyaline rings and
the length frequency distributions have been used to deter-
mine the age and growth rate of D. trunculus.
Introduction
Donacidae bivalves are some of the dominant organisms
capable of inhabiting the high energy environment of ex-
posed sandy beaches (Ansell 1983; Brown and McLach-
lan 1990), and as a consequence they are capable of reach-
ing high densities in some areas. The shape of the shell is
Communicated by O. Kinne, Oldendorf/Luhe
M, Ramdn ([]) R Abell6
Institut de Cihncies del Mar (C.S.I.C.), Plaga del Mar s/n,
08039 Barcelona, Spain
C. A. Richardson
School of Ocean Sciences, University of Wales-Bangor,
Menai Bridge, Gwynedd LL59 5EY, UK
9Springer-Verlag 1995
designed for allowing easy penetration into the sediment,
and these clams are excellent burrowers and thus ideally
adapted to life in the swash zone (Trueman and Ansell
1969; McLachlan and Young 1982).
The bivalve Donax trunculus is a warmwater temper-
ate species and is distributed throughout the Mediterranean
and the Atlantic from France to Senegal (Bayed and Guil-
lou 1985). In the western Mediterranean, D. trunculus oc-
curs in the surf zone down to depths of 2 m in the shallow
sublittoral, where it is the dominant macrobenthic organ-
ism in communities of generally low species diversity and
constitutes the target of a specific fishing activity carried
out by an artisanal fleet in that area.
Several studies on the growth of Donax trunculus have
been conducted, both in the Mediterranean and on the At-
lantic coast (Ansell and Bodoy 1979; Guillou and Le Moal
1980; Bayed and Guillou 1985; Maz6 and Laborda 1988;
Neuberger-Cywiak et al. 1990). These workers analysed
the monthly progression of the mean size of each cohort
identified in size frequency distributions and showed that
there is a seasonal pattern to shell growth. Attempts were
also made to analyse the external growth rings present on
the shell surface, although they were found to be difficult
to interpret (Ansell 1983). Recent growth studies of bi-
valves from the Mediterranean region, eg. Chamelea gal-
lina (Ram6n and Richardson 1992), have analysed micro-
growth patterns present in acetate peel replicas of shell sec-
tions, and this has allowed the relationship between envi-
ronmental factors and shell growth to be investigated. The
present paper is an analysis of the growth of D. trunculus
in the Mediterranean using both length frequency distribu-
tions and shell patterns.
Materials and methods
Samples of Donax trunculus were collected subtidally at approxi-
mately monthly intervals between May 1988 and April 1990 from a
sandy beach off Callera, on the eastern coast of Spain. Samples were
collected from a depth of between 0.5 and 2.5 m using a specially
666
designed experimental dredge similar to that used by local fisher- %~ ~ Jun 89 1
men but incorporating a smaller steel mesh size bag (4.5x4.5 ram).
This dredge was used to study the size distribution of the smaller
size classes. The anterior-posterior length of each clam was meas-
ured to the nearest 0.1 mm with a digital sliding vernier caliper %t ~ Jul 89
and length-frequency distributions constructed for each monthly ~
sample.
The method of Bhattacharya included in the ELEFAN software
package (Gayanilo et al. 1988) was used to identify and separate the %~ ~ Jul 88 %~ ~ Aug 89-g~
different components in the polymodal length-frequency distribu- 570 I
tions. Once identified, they were attributed to different cohorts of the
population.
In order to analyse shell growth of Donax trunculus, microgrowth %t ~ ~ Sep 88 Oct 8~
patterns present within the shell were analysed using the methodol-
ogy described by Ram6n and Richardson (1992). Subsamples of ten
shells of a range of sizes (17 to 36 mm in length) were taken from
each monthly sample and the right shell valve embedded in resin and t ~ oct 8B) ~ ~ No~
sectioned from the umbo to the ventral margin along the axis of max-
imum growth. Acetate peel replicas of the sectioned, ground, pol-
ished and etched surfaces were then prepared (Richardson et al. 1979;
Kennish et at. 1980). Acetate peels were examined in a transmitted
light-microscope. n =573
On first examination the shells of Donax trunculus did not ap-
pear [o have any marked external surface rings. However, a detailed
n28 i
examination of each shell, hacklit by a strong light source, revealed %t ~ Feb 89 %LC ~ Jan 90]
the occurrence of translucent or hyaline rings.
The yon Bertalanffy growth model parameters were estimated i 2.01 , ,~
from: (1) the size-age keys, obtained from counting the hy aline bands
and the assignment of a date of birth, using the program FISHPARM % Apr 891 Feb 9oq
(Saila et al. 1988) which estimates the growth parameters L~, the
asymptotic length, K, the yon Bertalanffy growth constant, and t0,
time of recruitment into the population; and from (2) an analysis of
the length-frequency distributions using the software package ELE- %I May 89
FAN (Gayanilo et al. 1988), which estimates the growth parameters n=170
L~ and K.
5 15 25 35 5 15 25 35
Length (mm) Length (mm)

Fig. 1 Donax trunculus. Monthly length-frequency distributions

Results from specimens collected between May 1988 and April 1990. Each
scale unit on the Y-axis is equivalent to 5% of population
A n a l y s i s of l e n g t h - f r e q u e n c y distributions

Fig. 1 shows the l e n g t h - f r e q u e n c y distributions of D o n a x

trunculus f r o m the m o n t h l y s a m p l e s c o l l e c t e d b e t w e e n 35/
M a y 1988 and A p r i l 1990. The p o p u l a t i o n s h o w e d two pe- []
0
riods of r e c r u i t m e n t each year, one during July to S e p t e m - 30
[] []
ber, with clams reaching a size o f 9 to 11 ram, and another ~ ] <>
b e t w e e n D e c e m b e r and February, with a size range of 8.5 ~ 2s ] D
[] []

to t 2 ram. T h e increase in size o f the recruited c l a m s be- ~ z0

t w e e n 1988 and 1990, d e t e r m i n e d f r o m the size p r o g r e s -
sion o f each cohort, is shown in Fig. 2. The figure suggests -~ i s 9 D
that b e t w e e n 1988 and 1990 the growth rate o f the winter
9 +
[] <> §
and s u m m e r recruiting cohorts was different. A f t e r an in- 10 [] <> 0 +
itial slow g r o w t h rate, clams w h i c h recruited to the p o p u -
lation during the winter o f 1987 and 1988 (W87, W 8 8 ) dis- s - I

p l a y e d a halt in g r o w t h around 20 m m b e t w e e n July and

1988 1989 1990
O c t o b e r 1988 and 1989, respectively, f o l l o w e d b y a rapid
Month
increase in g r o w t h in N o v e m b e r . H o w e v e r , those that were
lAW87 [] s 88 ow88 ~ s 89 + W89[
recruited during the s u m m e r s of 1988 and 1989 ($88, $89)
s h o w e d an a l m o s t constant i n c r e a s e in shell length during
both winter and summer. A n analysis o f the length-fre- Fig. 2 Donax trunculus. Patterns of shell growth of five cohorts be-
q u e n c y distributions, therefore, suggests that the p o p u l a - tween May 1988 and April 1990 determined from monthly progres-
tion contains two or three distinct cohorts, d e p e n d i n g on sion of mean size of each cohort in length frequency distributions.
the time o f y e a r when the s a m p l e is collected. (S summer; W winter recruited cohorts)

Fig. 3 Donax trunculus. Photomicrographs of acetate peel replicas
of shell sections. A Clearly defined (large arrows) and poorly de-
fined (small arrows) growth bands along the outer shell layer. B In-
ner shell layer with numerous growth bands (arrowheads). C Umbo
region with growth bands (arrows). D Alternating pattern of widely
spaced (large arrows) and narrow spaced (small arrows) growth in-
crements separated by growth bands. Scale bars=200 btm
Microgrowth patterns in the shell
The shell of Donax trunculus is composed of three layers,
an outer composite prismatic layer, a middle crossed la-
mellar layer, and an inner homogeneous/complex crossed
lamellar layer (Taylor et al. 1973). The clarity and inter-
pretation of the microgrowth patterns in acetate peel rep-
licas from these species were found to be more complex
than in the bivalve Chamelea gallina (Ram6n and Rich-
ardson 1992), which inhabits the same beaches although
at slightly greater depths than D. trunculus. In some re-
gions of the shell ofD. trunculus, microgrowth bands could
be clearly distinguished, whilst in others they were hardly
discernible (Fig. 3A). Bands were also observed in the in-
ner shell layer and in the umbo region (Fig. 3B, and C, re-
spectively), although in general they were less conspicu-
ous than the bands in the outer layer of the shell.
Examination of acetate peels of shell sections revealed
the presence in the outer shell layer of a pattern of widely
667
spaced growth increments alternating with a group of nar-
row increments (Fig. 3D). Separating each increment there
is a narrow growth band. In some regions of the shell the
bands were so closely spaced that they had a distinctive
appearance and were clearly marked with a slight dark
colouration which allowed the band to be traced through
the inner layer of the shell to the umbo (Fig. 4A and Fig.
3C, respectively). The dark structure in the outer layer (Fig.
4A) was often associated with a change in the direction of
shell growth (Fig. 4B) and was usually not associated with
an irregularity on the outer shell surface (Fig. 4C). Al-
though occassionally a small cleft in the shell surface did
coincide with the dark internal growth line (Fig. 4D).
Back lit by a strong light source the shell of Donax trun-
cuIus appears all but opaque except for the presence of
translucent or hyaline rings (Fig. 5 A-C). When the posi-
tion of the hyaline ring was compared with the internal
growth patterns, it was observed that the ring coincided
with the start of deposition of the internal dark band. The
hyaline ring was seen to be formed at the shell margin in
the summer (Fig. 5A). During the autumn and winter the
distance between the ring and shell edge steadily increased
(Fig. 5B, and C). The percentage of shells in monthly sam-
ples (June 1989 to June 1990) displaying a hyaline ring at
the shell margin is shown in Fig. 6. The highest percent-
age of shells with a ring at the edge were found in samples
collected between June and August, when the water tem-
peratures were at their highest (Fig. 6). Those shells
smaller than 15 mm did not display a hyaline ring, whereas
668
Fig. 4 Donax trunculus. Photomicrographs of acetate peel replicas
of shell sections. A Detail of thick dark structure in outer layer re-
lated to hyaline ring. Annual band can also be followed through mid-
dle layer (arrow). B Clearly defined dark structure associated with
change in the direction of shell growth (arrow). C Narrowing of the
growth increments (arrow) without any obvious cleft in outer shell
layer. D Cleft on shell surface (arrow). Scale bars in A=150 Barn;in
B=200 gm; in C and D=250 lain
all those larger than 25 m m displayed it (Fig. 7). In
view of the annual nature of the hyaline ring, these
structures were used to estimate the age of the shells of
this bivalve rather than the less easily distinguishable inter-
nal microgrowth patterns, which involved the time
consuming method of preparing acetate peels of shell sec-
tions.
Estimation of growth parameters
The von Bertalanffy growth parameters (K, L~) estimated
using data from measurements of the hyaline growth rings
and from the length frequency distributions showed simi-
lar values (Table 1). These values compare favourably with
those obtained in other studies of Donax trunculus from
the Atlantic coast of Europe and from the Mediterranean
(Table 1).
Discussion and conclusions
Microgrowth bands separated by growth increments of var-
iable width were observed in acetate peel replicas of shell
sections of Donax trunculus. The interpretation of the pat-
terns in these shells was more difficult than those previ-
ously observed in other species of subtidal bivalves, such
as Mercenaria mercenaria (Richardson and Walker 1991)
and Chamelea gallina (Ram6n and Richardson 1992). Both
micro- and macroscopic examination of the shells revealed
the presence of a prominent hyaline ring which was found
to be deposited in the summer between June and August.
Hyaline ring formation coincided with the deposition of
narrowly spaced growth bands during the summer months
when water temperatures were at a maximum of 22 to 27 ~
Ring formation, however, did not occur in clams smaller
than 15 mm, whereas those larger than 25 m m all displayed
a ring. Spawning in D. trunculus mainly occurs during the
summer months whilst the size at first maturation has been
established at a size of around 12.5 m m (Ramdn 1993).
During the summer the final stages of gonad maturation
take place with concomitant spawning, and this potentially
physiological demanding process presumably affects shell
formation and lead to the deposition of a hyaline ring. The
absence of a ring in sexually immature individuals smaller
than 15 m m lends support to such an interpretation.
An estimation of the growth rate of Donax trunculus
from an analysis of the length-frequency distributions
 669

100 30

E
80
25

'~ 60
o

-20
e~
40 E

5-_ 15
_= 20

o~
0 -10
J J A S O N D J F M A M J
1 989 1 990

Fig. 6 Donax trunculus. Monthly variation in percentage of shells

displaying hyaline ring at shell margin and, for comparison, water
temperature at time when each sample was collected (June 1989 to
June t990)

._.q
100 -mr t unto- mn

ii 1

80
iili
60 :ill iii

9 i!!i

40 ilii ii ~~ ~1
ii!i ii ::1
" ~iit
iiii
iiii
]iii
20 ii![

to
i !i
ii!ii , l!

10 12 14 16 18 20 22 24 26 28 30 32 34
Shell length (mm)
Fig. 5 Donax trunculus. Position of translucent hyaline rings (ar-
tvwheads) on surface of shells of different sizes backlit by a strong Fig. 7 Donax truncuIus. Relationship between occurrence of hya-
light source. A Shells collected in July 1989. B Shells collected in line rings at shell margin and shell size of clams collected between
October 1989. C Shells collected in January 1990. Scale bars=l cm June and August 1990

s h o w e d that there w e r e t w o recruitments o f clams to the
population each year, One cohort was recruited during the
winter whilst the other entered the population during the
summer. Although spawning takes place during summer,
the season is long enough to s h o w s o m e intraseasonal var-
iability in the gametes e m i s s i o n . Then, the s u m m e r cohort
may c o m e f r o m the first spawns (June-July), whereas the
winter cohort m a y c o m e from the latest spawns (Au-
gust-September). The growth rate of those that were re-
cruited during the winter s l o w e d d o w n and stopped the fol-
l o w i n g summer, w h e n they had reached a size of ca.
20 m m and had spawned. H o w e v e r , clams that were re-
cruited during the s u m m e r did not display in the f o l l o w i n g
s u m m e r the growth cessation observed during s u m m e r in
the winter recruited cohort. The s u m m e r recruited clams
did, however, s h o w a small growth cessation in April and
670
Table 1 Donax truncuIus. Es-
timates of the von Bertalanffy Area
growth parameters obtained by
various authors Mediterranean (Spain)
Mediterranean (Spain)
Atlantic (France)
Atlantic (France)
Atlantic (France)
Mediterranean (France)
Atlantic (Spain)
Atlantic (Spain)
a Age estimated from hyaline growth rings
b Age estimated from length-frequency distributions
c Age estimated from external growth rings
May ($88), June and July ($88) and December and Janu-
ary ($89) (Fig. 2) when they had reached a size of 24, 26
and 23 mm, respectively. There undoubtedly is a differ-
ence in the pattern of growth of the summer and winter re-
cruits during the summer. Since similar patterns of growth
were observed during two consecutive years, it is unlikely
that sampling error is envolved. The difference may be at-
tributed to a difference in the size of the clams and the time
of spawning.
The occurrence of a bimodal recruitment in D o n a x trun-
culus appears to be characteristic of the Mediterranean
populations (Moueza and Frenkiel-Renault 1973; Bodoy
and Mass6 1979; Bayed and Guillou 1985), whereas in the
Atlantic populations there is always a single annual recruit-
ment (Ansell and Lagard~re 1980; Guillou and Le Moal
1980).
Our observations on shell growth in D o n a x trunculus
from the Mediterranean are different from those obtained
for D. trunculus from the Atlantic coast of Europe. Stud-
ies carried out by other authors on the seasonality of shell
growth in D. trunculus have shown that the period of
growth cessation occurs during the winter rather than in
the summer months (Ansell and Lagard~re 1980; Guillou
and Le Moal 1980; Bayed and Guillou 1985). However,
Ansell and Bodoy (1979) suggest that in addition to a de-
crease of growth in winter, another one occurs in the sum-
mer. The Atlantic populations of D. trunculus occur inter-
tidally and are exposed to lower seawater temperatures
than the subtidal Mediterranean populations, where water
temperatures rarely fall below 12~ The lower water tem-
peratures on the Atlantic coast probably cause the shell
growth of D. trunculus to slow down to a greater extent
than in the Mediterranean.
The present study has revealed the presence of hyaline
growth rings on the shell surface of D o n a x trunculus, and
these rings have been used successfully to estimate the age
of this bivalve. Traditionally, it had been considered that
this species did not form annual growth rings in the Med-
iterranean. Therefore, estimates of the growth of D. trun-
culus had relied entirely on the interpretation of length-fre-
quency distributions (Moueza 1972; Bodoy 1982; Neuber-
ger-Cywiak et al. 1990). In view of the difficulties encoun-
tered in establishing age-length relationship, von Berta-
L~ K to Source
41.8 a 0.71 -0.354 Present study
46.0 b 0.58 --0.416 Present study
48.90 b 0.38 0.29 Guillou and Le Moal (1980)
35.55 b,c 0.785 - Ansell and Lagard~re (1980)
32.25 b.c 0.678 - Ansell and Lagard~re (1980)
35.99 b 0.956 0.699 Bodoy (1982)
43.80 b 0.97 0.10 Fernfindez et al. (1984)
52.84 b 0.55 - Maz6 and Laborda (1988)
lanffy growth parameters have usually only been estimated
for the Atlantic populations (Table 1). This work along with
the earlier work of Bodoy (1982) is the first attempt to es-
timate growth parameters for the Mediterranean popula-
tion. Nevertheless, other studies in the Mediterranean have
referred to the seasonality of shell growth but have not
sought to establish relationships between length and age
(Ansell and Bodoy 1979; Bodoy and Mass6 1979). Whilst
our estimates of K and L~ fall within the limits of those ob-
tained for the Atlantic populations by other authors (Table
1), the value of K determined by Bodoy (1982) is higher
and the value of L= is lower than our estimates for the Med-
iterranean. The growth rate of D o n a x trunculus is higher
(K=0.6 to 0.7) than that of Chamelea gallina (K=0.3 to 0.4;
Ramdn 1993), another bivalve which inhabits the same
beaches. This is not surprising since D. trunculus, which
lives in the more unstable environment of the surf zone,
has a short life-span (maximum 2 to 3 yr), (Moueza 1972;
Bayed and Guillou 1985; Maz6 and Laborda 1988; present
results) compared to C. gallina (3 to 4 yr; Ramdn and
Richardson 1992; Ram6n 1993). The longevity of D. trun-
culus in the Mediterranean is also shorter than in Atlantic
populations which live to about 5 yr (Ansell and Lagard~re
1980; Guillou and Le Moal 1980).
Acknowledgements Our gratitude is expressed to D. J. Lleonart
for his encouragement and statistical advice. Financial support was
provided by the Generalitat de Valencia.
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