South African Journal of Marine Science

ISSN: 0257-7615 (Print) (Online) Journal homepage: https://www.tandfonline.com/loi/tams19

Changes in the structure of a shallow sandy-beach

community in Peru during an El Niño event

W. E. Arntz, T. Brey, J. Tarazona & A. Robles

To cite this article: W. E. Arntz, T. Brey, J. Tarazona & A. Robles (1987) Changes in the structure
of a shallow sandy-beach community in Peru during an El Niño event, South African Journal of
Marine Science, 5:1, 645-658, DOI: 10.2989/025776187784522504

To link to this article: https://doi.org/10.2989/025776187784522504

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The Benguela and Comparable Ecosystems
Payne, A. I. L., Gulland, J. A. and K. H. Brink (Eds). S. Afr. J. mar. Sci. 5: 645-658
1987 645

CHANGES IN THE STRUCTURE OF A SHALLOW SANDY-BEACH

COMMUNITY IN PERU DURING AN EL NINO EVENT

W. E. ARNTZ·, T. BREyt, J. TA RA ZONA •• AND A. ROBLESt

Many infaunal sandy-bottom communities in shallow waters of the Peruvian upwelling system are inhabited
by large coexisting populations of the surf clams Mesodesma donacium and Donax peruvianus as well as by
the anomuran mole crab Emerila ana/oga. Under normal conditions, equilibrium states are possible with any
one of these species dominating. A Mesodesma community south of Lima in Peru was investigated over 2,5
years, covering periods prior to, during and after the EI Nino (EN) of 1982-83. It was revisited several times
later. Growth, recruitment and mortality and, therefore. production of Mesodesma and Donaxvaried to some
extent before EN. However, during the event Mesodesma became locally extinct and had not recoloniz.ed the
area by July 1986, three years after the return of normal temperatures. Donax, which took over immediately
after EN, never reached the densities of the former dominant Mesodesma. Emerila remained a rare species as
well, whereas spionid polychaetes increased in importance. The medium-term effect of the exceptionally strong
EN of 1982-83 appears to have been very marked on Peruvian sandy beaches. A comparison is made with
other shallow-water communities of the upwelling system, and the importance of EN in terms of oscillations of
clam stocks off Peru and Chile is discussed.

In baie infauna-gemeenskappe van sandbodems in die vlak water van die Peruaanse opwelstelselleef groot
bevolkings van die sandmossels Mesodesma donacium en Donax peruvianus saam, sowel as die molkrap
(Anomura) Emeri/a ana/oga. Onder gewone omstandighede is ewewigstoestande moontlik waarin enigeen van
die spesies kan oorheers. 'n Gemeenskap van Mesodesma is vir 2,5 jaar suid van Lima in Peru ondersoek, voor,
tydens en na die El Nino (EN) van 1982-83. Oit is later verskeie kere herbesoek. Groei, rekrutering en
mortaliteit en dus ook produksie van Mesodesma en Donax het voor EN ietwat gewissel. Tydens die episode
het Mesodesma egter plaaslik uitgesterf en het teen Julie 1986 nog nie die gebied herkoloniseer nie, drie jaar na
die temperatuur tot die normale teruggekeer het. Donax. wat dadelik na EN die oorhand gekry het, het nooit
die digthede van die voorheen oorheersende Mesodesma behaal nie. Emerila het ook skaars gebly, maar
spioniede polichete het belangriker geword. Op die medium termyn blyk dit datdie uitsonderlik sterk EN van
1982- 83 'n baie opmerklike uitwerking op Peruaanse sandstrande gehad het. 'n Vergelyking word getref met
ander vlakwatergemeenskappe van die opwelstelsel, en die belangrikheid van EN wat skommelings in die
sandmosselstapels langs Peru en Chili betref, word bespreek.

Macrobenthic infaunal communities of intertidal
and shallow-water sandy bottoms of the Peruvian
upwelling system have little in common with other
tropical, benthic communities in low latitudes. With
a low number of species, some of which are highly
dominant, and a high biomass, they rather resemble
the benthic communities of temperate areas (Arntz
and Rumohr 1982, Brey 1984, Moller et al. 1985),
which also have similar temperatures in summer. On
the other hand, seasonal temperature variation does
not play such an important role as it does in
temperate marine areas (Rachor and Gerlach 1978,
Arntz and Rumohr 1982, 1986) because the winter
values are much higher in the upwelling region and,
therefore, the range of temperature changes is
reduced. Temperate marine areas suffer mainly from
occasional catastrophes such as ice winters (e.g.
Crisp 1964, Ziegelmeier 1964, 1970), oxygen defi-
ciency due to exceptionally warm summers (e.g.
Weigelt and Rumohr 1986) and, recently, eutrophi-
cation (Rosenberg et al. 1986). In contrast, the
upwelling area off Peru and northern Chile is struck
every 5-7 years by EI Nino (ENSO) events which
affect the fauna both in positive and negative ways
(Arntz 1986).
This report deals with the shallow-water sandy
bottom community of Santa Maria del Mar, about
60 km south of Lima (l2°S), especially with two of
the three dominant species living in this community
under normal conditions. Density and length fre-
quency composition of the populations of the migra-
tory surf clams Mesodesma donacium and Donax

* Alfred-Wegener-Institut fur Polar- und Meeresforschung, 2850 Bremerhaven, F.R. Germany

·t Institut fUr Meereskunde, 2300 Kiel, F.R. Germany
** Universidad Nacional Mayor de San Marcos, Lima 100, Peru
t E. de Habich 246, Urb. Ingenieria, Lima 31, Peru
· Contribution No. 34 of the Alfred-Wegener-Institut
· Manuscript received: September 1986
 The Benguela and Comparable &osystems
646 South African Journal of Marine Science 5
peruvianus were registered before, during and after
the exceptionally strong EI Nino of ]982-83, reveal-
ing interesting changes in the populations of these
species and the community under consideration. The
results allow for some conclusions to be drawn as to
the stability of this type of upwelling community.
The third important species, the mole crab Emerita
analoga, has previously been investigated in detail by
Penchaszadeh (] 97]); during the time of the present
investigation it was only of minor importance in
Santa Maria.
The present study was originally initiated to investi-
gate variations in density, growth and mortality of
the two bivalve species involved. Therefore, addi-
tional information on other, less important co-
existing species or groups was restricted to density
values and is not analysed in great detail here (but see
Tarazona et al. 1985, in prep. a, b).
MATERIAL AND METHODS
In Santa Maria del Mar, samples were taken at
intervals of between 2 weeks and 2 months at two
sites (Station I= =
southern part, Station 2 northern
part of the bay) about 300 m apart. The sediment of
the two stations showed no apparent differences in
grain size (70-80 per cent > I25 ~m) or organic
content (1,5-2,5 per cent). At Station I, quantitative
sampling started in Decem ber 1980 (two years before
EN) and finished in November 1983 (five months
after EN). For the five months after EN, quantitative
samples are also available from Station 2. Data on
length composition of the two bivalve populations
are available for the full times pan of three years
including EN; from June 1981 there is also informa-
tion on the numbers of Emerita analoga and other
macro benthic fauna.
Sampling was carried out between the shoreline
and the breaker zone, during low tide, at a water
depth of 50 ± 20 cm. During and after EI Nino,
when the M. donacium stock was killed off in
shallow water, additional samples were taken in
deeper water down to 10 m. At Station I, ]0 replicate
samples were taken on each sampling date by means
of a push-box with a surface area of 200 cm2 (see
Rumohr and Arntz 1982) that penetrated 20 cm into
the sediment. Unlike Mesodesma mactroides which,
according to Defeo (1985a), may befound as deep as
40 cm in the sediment, M. donacium was never
found more than 10 cm deep during this study. At
Station 2, a collecting bag attached to a semi-circular
wooden frame (40 cm wide, 20 cm high) was used: on
1987
each date, the bivalves were dug out by hand over
half an hour and washed into the bag by the receding
waves. This gear is used by the Peruvian clam
fishermen in shallow water, but with a much larger
mesh size. The mesh used for this study was I mm.
The samples taken at Station I represent quanti-
tative density values whereas those of Station 2
cannot be referred to a certain area but, because of
the manner of sampling, are representative of the size
composition of the two bivalve populations.
The bivalves were preserved in 5-per-cent formalin
and measured to the nearest millimetre below. For
estimation of growth, two methods were used. First,
a simple linear regression was drawn of median
cohort length at time t, against time t, which pro-
vided an average length increment per unit of time
for the whole lifespan of the cohort. Second, a micro-
computer-supported analysis of length frequency
samples named ELEFAN was used. This was origi-
nally developed by Pauly and his associates for the
investigation of tropical fish stocks, and was adapted
for benthic invertebrates from temperate latitudes by
Brey (1986) and Brey and Pauly (1986). In contrast to
traditional fish stock assessment, the individual age
of animals need not be known. A seasonally oscil-
lating von Bertalanffy growth curve (Pauly and
Gaschiitz 1979) was fitted to the length frequency
data of each single cohort, which were regrouped in
5-mm (Mesodesma) and 2-mm (Donax)
classes in an objective and reproducible way (Pauly
and David 1981):
Lt = Lx, (I - e-K [(t - to) + 2~ sin (21T (t - ts))]) ,
where Loo is the asymptotic length, K is the growth
constant, C is a constant expressing the amplitude of
the growth oscillation, ts is the starting point of
=
oscillation with respect to t 0, to is the age at zero
length and L, is the length at age t.
The ELEF AN method does not yield exact con-
fidence limits for the growth parameters, and the
range of growth-parameter combinations with simi-
lar good fits depends mainly on the sample structure.
In the results (Tables II and IV), most of the different
combinations of K and Loo which yield identical
maximum goodness of fit have been included as a
simple measure of dispersio!!.
Production and the PI B ratio of Mesodesma
donacium were calculated for the period December
1980-July 1982 (Cohort 2) by means of the increment
summation method of Crisp (1984). Mean individ ual
weights and biomasses were calculated from length-
weight relationships, which have been established for
/987 Arntz et aJ.: Beach Community Structure Changes in Peru during E/ Nino 647

different months ofthe year. To determine shell-free lal

dry weight (SFDW), the soft parts were dried at a 22
temperature of 60°C until the weight remained 20
constant.
18
An estimate of annual mortality in the Mesodesrna 0'::
donacium population was made by fitting the single w 16
negative exponential mortality model to the data '"
:)

'<
used for calculating production:
'"
%w
lbl
, .
....• /'-'
24
•... 22 .pc...., t· \
Nt = Noe-Zt . // I \. /! Long-term "
20 . I ~.. average •
where No is the number of bivalves in a given cohort \,._.... 1973-1982
18
born at time zero, Nt is the number of bivalves in the
same cohort alive at time t and Z is the annual J A SONDJ FMAMJ J ASONDJ FMAMJ J A SO
instantaneous rate of total mortality. 1981 1982 26 /\ lei
24/V\
RESULTS

Sea surface temperatures before and during EI Nino

22

20

Sea surface temperatures (SST) are available for
Callao, 60 km north of Santa Maria, and Pisco, 160
km south, both for the long-term mean and the
period January 1981-December 1983, the period
that includes the EN of 1982-83 (Fig. I). SSTs were
below the long-term mean in 1981 and the first half of
1982, but were much higher than normal during EN.
For Santa Maria, measurements are available only
for the period December 1982-0ctober 1983; it can
be assumed, however, that the positive anomalies at
this site during EN were similar to those observed at
the other localities.
Density and dominance of Mesodesma donacium
Densities of Mesodesrna donaciurn were highest in
mid-1981 when 9160 individuals·m-2 were found at
Station I (Fig. 2); their mean length was 32 mm. At
this time only one age-class (Cohort 2, recruited in
December 1980) was present (see Fig. 5). Although a
new cohort recruited in February/March 1982
(Cohort 3), population numbers declined continu-
ously until June 1982, when there were only 885
individuals·m-2 with modal lengths of 10 mm
(Cohort 3) and 50 mm (Cohort 2). Another recruit-
ment (Cohort 4, October 1982) then produced an
increase until December 1982/January 1983, with
densities of 2 940 and 3 190 individuals·m-2 respec-
tively. The modal lengths of the three different
cohorts were 15, 32 and 54 mm. In February 1983
18
\ •
\.........
16
JFMAMJJASO
1983
Fig. 1: Sea surface temperatures at (a) Callao (12°5),
(b) Pisco (14°5) and (c) temperatures measured
during EN at Santa Maria. Callao and Pisco data
from the Peruvian Navy. Callao; Santa Maria data
measured at 1 m depth during high tide
density declined to I 595 individuals·m-2• Up to this
month, with three exceptions at the very start of the
quantitative sampling, when it was "only" between
86,4 and 91,5 per cent because of a temporary
increase of Semimyti/us aJgosus (see Figs 4 and 9a),
Mesodesma had always contributed> 95 per cent of
the animals. However, in February 1983, 17,2 per
cent of the bivalves were no longer able to close their
valves.
From March 1983, no more live M. donacium
were found in shallow water either at Station I or 2.
Regular sampling until November 1983 and occa-
sional checks thereafter showed no recovery up to the
time of writing (July 1986). Diving in deeper water
revealed a few patches with densities between. 175
and 360·m-2 at 6 m and 265 individuals·m-2 at 10 m in
May 1983; however, from June 1983 these had
disappeared as well. Comparative sampling in the
Department of Lima and farther south showed that
M. donacium stocks survived EN 1982-83 only
south of 1405. Commercial exploitation almost
 The Benguela and Comparable Ecosystems
648 South African Journal of Marine Science 5 1987

10000 ,

I
I

Mean and standard error i

m~//:./-;~·:,;~"-"
//~
~
w
~
8000
• ~;:a:":~':bk>1f/~L/~'~5?}?~:~///.
w ,
'"
~ 6000
f I

~
~
:'l
Q
:>
15 4000
~
0
'"w.., 2000 •
§ I
•I
I
I
I
I No M. donacium found
0 I

DJ FM'A.MJ JjASONDJ FMAMJ JASONDJ FM'AM'J,JIA!S'ON,D

1981 1982 --------- 1983 I

Fig. 2: Densities (number.m-2) of Mesodesma donacium at Santa Maria, Station 1, December 1980-November
1983

,
,
500
I
Mean and standard error I
I
I
I

f~L'.~!~<?: 1~82,-;8i/'l

300

I I
200

I
t
I
I
I
I
I I
100 I I
I I
I

tft I
I 1 i
I if
I

0
h f f f ! £ t f:f t •
D J F M A M J JASONDJ F M A M J J A SON D J M A M J J A SON D
1981 I 1982 I 1983 I
Fig,3: Densities (number.m-2) of Donax peruvianus at Santa Maria. Station 1. June 1981-November 1983
1987 Arntz et al.: Beach Community
Mesodesmo
I
~ ,
'" 80
III
~ \
~ f:IJ
>-
~ 40
i 20
JUN. '81 DEC. '81 JUN. '82
Fig. 4: Percentage contribution of different species/groups of species to the total numbers at Santa Maria,
Station 1, June 1981-November 1983
collapsed in Peru during and after EN and had not
recovered by July 1986.
Density and dominance of Donax peruvianus
Donax peruvian us never reached as high a density
at Station I as did M. donacium. Even after the latter
species disappeared there, Donax densities remained
comparatively low. Throughout the investigation,
densities oscillated between 0 and 60·m-2 with the
exception of the first sampling in June 1981, when it
was 170.m-2, and dominance values always remained
low (0,1-5,7 per cent, see Fig. 4) up to April 1983.
Then, despite low densities, it became the dominant
species with percentages between 59,4 and 100 per
cent. In October/November 1983, densities were
between 183 and 185 individuals·m-2 (Fig. 3).
At Station 2, Donax increased from a low density
of 13,5.m-2 in July 1983, when the EN had just
finished, to values of 133,5 in October 1983. At this
station, the Mesodesma: Donax ratio was much
more even between February 1982 and February
1983.
Densities of Emerita ana/oga and other macrobenthos
At Station I, densities of Emerita analoga were
extremely low throufh<;)Ut the invest~gation, va.rying
between 0 and lO·m- with the exception of Apnl and
November 1983, when 85·m-2 (mostly juveniles) were
found. No specimens at all were recorded between
March and October of 1982 and June and October of
Structure Changes in Peru during EI Nino 649
,.- - "'\
, (,oonox
\
, \
,
," \
\
• ,I ~
DEC. '82 JUN.'83
1983. Dominance of E. analoga always remained 1 <
per cent up to February 1983, but in April and May
1983 it contributed about one-third of the animals,
and in November 28,5 per cent (see Fig. 4).
The density of other macrobenthic organisms
hardly ever exceeded 100·m-2 at Station 1. Notable
exceptions were October / November 1981, when the
mussel Semimytilus algosus used M. donacium as a
secondary hard bottom, attaining densities of 540
and 363.m-2 respectively, and in October there were,
in addition, 107 polychaetes.m-2 of various species.
During the EN, invasions of macrobenthos became
more frequent: 155young stalked barnacles Pollicipes
elegans and many hermit crabs Pagurus perlatus
appeared in April 1983, and 38 isopods (Excirolana
brasiliensis).m-2 were recorded in November 1983.
After November 1983, spionid polychaetes of the
genera Dispio and Scololepis became members of
the community at Station 1 (Tarazona, unpublished
data). Generally, other macrobenthic species were of
minor importance in this community before EN but
increased during the event. This conclusion is also
reflected by the dominance values (Fig. 4).
At Station 2, E. analogacontributed 10,8-91,7 per
cent between October 1982 and April 1983. In March
and April 1983, there was some shift of dominance
between D. peruvianus and E. analoga, and then
Donax took over until November 1983, when
Emerita took first place again as a result of recruit-
ment. At this station, other macrobenthos never
contributed more than 7,2 per cent except in Novem-
ber 1983, when the isopod Excirolana brasiliensis
invaded the area, as it did at Station 1, and raised the
share of this group to 31,4 per cent.
 The Benguela and Comparable Ecosystems
650 South African Journal of Marine Science 5 1987

Sarrpling Sampling
date date
(1981) (1982)
10
8.1. 10
15.1.
10 26.1.
Sampling 18.2.
date
(1980) 10 15. 3.
10 2.4
10 7.4. 10
20.4. 17. 4. 17. 4
10 3.5. 29. 4.
11.5.
10 10
31.5. 10 1.6.
10 116.
21.6.
§ 10
2.7.
10
>- 20.7. 10
28.7. 10
~ 10
11.8. 10
30.7.

§ 10
1.9. 10
23.8. 10
5.\1
ll: 10 13.9.
28. 9. 10
3.10 10
IG1O. 8.1Q
10 2Q1Q 1\110.
27.10. JOKl
4.11.
10
23.11. 10
11.12.10 l5.12.
10 10
31.12. 31.12. Xl 20 30 40 50 60 70 80
10 20 30 40 50 60 70 80 10 20 30 40 50 60 70 80
LENGTH (mm)

Fig. 5: Length frequency distribution and growth curves of different cohorts of Mesodesma donacium at
Santa Maria, Stations 1 and 2, 1980-1982. The two months when M. donacium was found in 1983 are
omitted because there was no growth

Recruitment and growth of M. donacium
The mean length increments, derived from linear
regressions of length against time, are shown in
Table I.
The growth curves described in this section are
shown in Figure 5 and the corresponding growth
parameters in Table II. During the whole period of
investigation (April I980-March 1983) the growth of
three different cohorts was observed.
Cohort I recruited in early 1980 (before sampling
started) and vanished by the middle of 1981. As
indicated by the mean length increment
(3,9 mm.month-I) and the growth parameters, this
cohort grew very fast, and the clams reached a mean
length of 70,0 mm after their first year of life. From
photographs taken in August 1980, it is evident that
densities were about as high as in 1981.
Cohort 2 recruited during December 1980-January
1981 and was still present at the end of 1981. Growth
was much slower in this cohort. The mean length
increment was only 1,7 mm·month-I and, during the
first year of life, a mean length of less than 50 mm
was reached. It was not possible to fit a single growth
curve through the whole lifespan of this cohort,
because there seems to have been a break in growth
in the middle of 1982. After this date, Cohort 2
showed almost no growth (see Fig. 5), and it became
completely extinct during 1983.
Cohort 3 recruited in early 1982 and showed a slow
growth rate similar to that of Cohort 2 during the
first half of the year. The mean length increment
amounted to 1,2 mm·month-'. AdditionaUy, this
cohort was mixed with an irregular recruitment in
November 1982, so the growth function was fitted
until September 1982 only.
Recruitment and growth of D. peruvianus
The mean length increments of this species are
given in Table III. The growth curves are shown in
Figure 6 and the corresponding growth parameters
are listed in Table IV.
For this species also, the growth of three different
cohorts has been analysed. Cohort I recruited in late
1979-early 1980 (before sampling started) and dis-
appeared in early 1981. The mean length increment
was 1,1 mm.month-landanaveragelengthof23 mm
was reached after one year.
Cohort 2 'recruited in December 1980 and was
present until early 1982. Cohort 3 recruited in early
1982 and disappeared during 1983. The rates of
growth of both these cohorts were slower than that of
Cohort I, the mean increments being 0,9 and
1,0 mm·month-I respectively. After the first year of
life, their average length was still less than 20 mm.
1987 Arntz et af.: Beach Community Structure Changes in Peru during Ef Nino 651

Table I: Length increments calculated from linear regressions of median length (mm) against time (days) for M. donacium

Cohort Period e.I ·day-' 95% range t:;1·month-1 n ,2

I Apr. 80-Jan. 81 0,128 ±O,O09 3,9 14 0,986

2 Dec. 80- Dec. 82 0,058 ±0,008 1,7 31 0,839
3 Jan. 82- Sept. 82 0,041 ±0,024 1,2 II 0,608

Table II: Growth parameter values of a seasonally oscillating von Bertalanffy growth function for M. donacium

Cohort Period K Leo Range C WP Index

1 Apr. 80-Feb. 81 1,13 124 0,98/134-1,28/114 0,91 0,64 0,819

2 Dec. 80-Jul. 82 0,38 110 0,35/117-0,41/107 0,84 0,63 0,581
3 Jan_ 82-Sep_ 82 0,32 84 0,20/104-0,44/ 64 0,84 0,63 0,793

Range == lower-upper limit of combinations of K and Leo values with maximum index (C and WP fixed)
Index = index for goodness of fit ~ 1,0
WP = winter point (WP = Is + 0,5)

Table III: Length increments calculated from linear regressions of median length (mm) against time (days) for
D. peruvianus

Cohort Period t:;1-dai' 95% range t:;1-month-' n ,2

1 Apr. 80-Jan_ 81 0,D35 ±O,O07 1,1 9 0,949

2 Mar. 81-Apr. 82 0,031 ±O,OI2 0,9 14 0,721
3 Feb. 82-Mar. 83 0,032 ±O,OIl 1,0 15 0,741

Table IV: Growth parameter values of a seasonally oscillating von Bertalanffy growth function for D. peruvian us

Cohort Period K Leo Range C WP Index

1 Apr_ 80 Jan. 81 1,00 46 0,52/58 1,46/34 1,03 0,67 0,973

2 Mar. 81 Apr. 82 1,17 35 1,15/35-1,18/35 0,98 0,66 0,620
3 Feb. 82-Mar_ 83 0,70 42 0,45/58-0,95/26 0,98 0,60 0,685

Range = lower-upper limit of combinations of K and Leo values with maximum index (C and WP fixed)
Index = index for goodness of fit ~ 1,0
WP = winter point (WP == Is +
0,5)

A fourth cohort, recruited in 1983, was not
analysed. However, the very slow growth of this
cohort during 1983 is clearly indicated in Figure 6.
Biomass, production, P(ii ratio and mortality of
M. donacium
These parameters were calculated for Cohort 2
during December 1980-December 1982 from the
samples taken at Station I. The length-weight relation-
ships used for the calculation of mean individual
weights and biomass are given in Table V.
All biomass figures presented here refer to shell-
free dry weight (SFDW). They can be converted to
shell-free wet weight (SFWW) by the empirical
relationship SFWW = 4,726 . SFDW (n = 56,
standard error = 0,100).
Between December 1980 and December 1982, the
mean biomass amounted to I 100 g SFDW.m-2, the
lowest value observed being 275 g SFDW·m-2 and
the highest I 996 g SFDW.m-2•
During the same period, total production of
Cohort 2, which is nearly the total production of the
whole population during that time, amounted to
2900 g SFDW.m-2• This is equivalent to a production
of 13,7 kg SFWW·m-2• As indicated by Figure 7,
cumulative production increased linearly from
January 198 I until the middle of 1982, but then it
ceased for the remainder of the year.
 The Benguela and Comparable Ecosystems
652 South African Journal of Marine Science 5 1987

Sarrpling
Sampling date
dote (1983)
(1982) 4. 1.
Sampling 18-I.
dote
(]981) 20
112.
18.2.
Sampling 25. 2.
20
dote
14.1 16. 3.
(1980) 21. 3.
20 20
2.4.
7. 4. 74.
17 4. 0
20. 4. 20 20.4.
24.4.
20 3.5.
11.5.
20 20
31. 5. 20 I. 6. 2.6.
20 13.6.
21. 6. 20
2.7 306.
~ 20
20.7 20 20
>-
u 20 28.7. 307 26.7
zw
g 11.8- 20
218.
'"...
20
20 29. 8.
5.9.
20 13.9.
20
28.9. 20
110.
20 8-10
20.1Q 19.10 22.10.
20
4. n. 3010. 10 20 30

20
20
11.12.
20 10 20 30 15.12.
10 20 30
31. 12.
10 20 30
LENGTH Imm)

Fig. 6: Length frequency distribution and growth curves of different cohorts of Donax peruvian us at Santa
Maria, Stations 1 and 2, 1980-1983

A PI Ii ratio of about 2,6 for the whole period of values for mean annual biomass (I 060 and UOO g
observation was calculated from these biomass and SFDW.m-2for 1981 and 1982) the annual PIB ratio
production figures. The annual production of Cohort is 1,8 for 1981 and 0,9 for 1982.
2 amounted to I 900 g SFDW·m-2 in 1981 and Mortality of the Mesodesma donacium population
I 000 g SFDW·m-2 in 1982. With the corresponding was analysed by following the decrease in numbers
over time. A plot of the natural logarithm of
abundance against time is given in Figure 8. Between
Table V: Length-weight-relationships for Mesodesma the middle of 1981 and the middle of 1982 the single
donacium
exponential mortality model seems to fit the data.
Total mortality Z, calculated from the regression
Period a b Number of Correlation
animals coefficient shown in Figure 8, amounted to 2,5.
March -5,344 3,116 35 0,967
May -5,249 3,013 36 0,944
June-July -5,227 2,995 38 0,945 DISCUSSION
October -4,260 2,390 27 0,900
November -4,794 2,731 29 0,908
December -5,010 2,812 24 0,949 Mesodesma community before EI Niiio
Log (W):= a + b . log (L). where L is the length (mm) and W is the
shell-free dry weight (g) Santa Maria beach forms part of the Peruvian
1987 Arntz et aJ.: Beach Community Structure Changes in Peru during EI Nino 653

w
e:w
~ _ 8000

:i! 3000 Z
« w
§ '"~ 4000
w
o '"«
~ 2000 o
I-
o o
:r:
l)
w
ffi
Q. o In N,- -10,502-2,539'
n. 14(10June 1981-July 1982)
o 0
o

3 '" 1000
III r·O,916 o
~ 1000 o :<
::> 1 Jon, 1981~, - 0
o o Z
w
w
500
'"
u..

.
I
-'
-' o , JAN.1."81 JAN.1.'82 JAN.l:83
I

~ JAN.1. '81 JAN,I. '82 JAN.l:83

Fig. 8: Mortality of Mesodesma donacium, Cohort 2,

Fig, 7: Cumulative production of Mesodesma donacium, Station 1. Santa Maria
Cohort 2. Station 1. Santa Maria

only, the Mesodesma population at Station I in

upwelling system, which has been described as a
cold-temperate ecosystem despite its being at a very
low latitude (Rosenberg et aJ. 1983). Under normal,
upwelling conditions, many shallow-water sandy
bottom communities in this area are characterized by
a small number of highly dominant species with high
densities and biomasses, fast growth and an extremely
rich production. In most cases either Mesodesma or
Donax and/ or Emerita are the main dominant, and
usually one or two of the other species coexist in the
same area although in reduced densities (Osorio et al.
1967, Penchaszadeh 1971, Defeo 1983, 1985a, b,
Tarazona et al. 1985).
On Peruvian beaches, Donax peruvian us and
Emerita analoga can also form extremely dense and
dominant populations in similar sh<;tllow-water sand
environments when M. donacium is absent or in low
numbers (Penchaszadeh 1971, Tarazonaetal. 1985).
Both maximum densities and biomasses of M.
donacium exceed the values presented by Penchas-
zadeh (op. cit.) for E. analoga in Peru, by Tarifeno
(1980) for M. donacium in Chile and by Defeo
(1985b) for M. mactroides in comparable systems on
the Atlantic coast of Uruguay.
In the pre-Nino year 1981, when temperatures
wer~ below the long-term mean, mortality Z and thus
PI B ratio were about 2,0 for the total population
(rather a temperate than a tropical value according to
Ansell et al. 1978). The Mesodesma production then
amounted tosome2 000gSFDW.m-2, whichcorres-
ponds to a wet weight (including shells) of 35,5
kg.m-2• This is an extraordinary value for a sandy
beach. From a calculation similar to that of Penchas-
zadeh (1971) who, however, referred to biomass
favourable years might have produced more than I
ton of surf clams per metre of beach. Apparently a
great part of this production is utilized by small-scale
fishermen and their families. At beaches where
densities of M. donacium are high, hundreds of
people can often be observed exploiting clams of a
size hardly exceeding 40 mm.
During normal times, the very dense M. donacium
populations at the two stations in Santa Maria left
few gaps in time or space for other macrobenthos. D.
peruvianus attained maximum densities of about
170·m-2 only at the start of the study, and E. analoga
never exceeded 10·m-2• The initially successful
attempt of the mussel Semimylilus algosus to colonize
the Mesodesma "hard bottom" (cf. Fig. 9a) with
densities up to 740·m-2 ended abruptly when the
clams were covered by sediment during a period of
strong swell. Other macro benthic species apparently
never had a real chance to settle.
In a relatively warm year such as 1980, the clams
grow to a length of 70 mm, whereas in colder years
they attain only 50 mm. This initial growth is much
faster than that recorded by Tarifeno (1980) for
M. donacium in Chile and by Defeo (l985b) for
M. mactroides in Uruguay, although densities in the
present case were higher. However, the values of Loo
calculated for the Santa Maria population (110-
124 mm) correspond well to the Loo of 115 mm
calculated by Tarifeno (op. cit.) for Chilean popula-
tions of surf clam. Clams of this size were never
found during the present investigation although they
appear occasionally on the market. Probably the
intense exploitation depletes a cohort a long time
before its members reach maximum size. Also, some
 The Benguela and Comparable Ecosystems
654 South African Journal of Marine Science 5
emigration of large animals to deeper water cannot
be totally excluded.
Conversely, the size frequency distributions and
especially the catch curve (Fig. 8) of Cohort 2 reveal
that only part of the population recruits in shallow
water among the adults, where the present sampling
took place. There was a 4,5-fold increase in abund-
ance of Cohort 2 from February 1981 (2 050 indivi-
duals.m-2) to June 1981 (9 150 individuals.m-2),
though the average length of the animals increased
from 16,5 to 31,5 mm during the same period. As
gear selection was negligible here due to the I-mm
screen used, the only explanation for this phenome-
non is that many juveniles recruit in deeper water and
migrate towards the shore later, increasing the num-
bers at the survey stations. It has proved impossible
to investigate this suspected seasonal migration in
more detail because it would have complicated
immensely the sampling procedure.
Ansell et al. (1972) had similar difficulties with
Donax populations in India and used a transect to
minimize the problem, as did Tarifeno (1980) in his
Mesodesma study. Interestingly, this suspected mi-
gratory behaviour would be the reverse of what has
been observed in Chilean M. donacium communities
(Tarifefio op. cit., M. Sanchez pers. comm.), in which
the juveniles recruit nearshore and the adulls emigrate
to deeper water. This leads to apparently slow
growth in shallow-water sites, according to the length
frequency distributions. With either behaviour, the
juveniles would avoid settling amongst the larger
specimens where they would suffer intense inter-
actions between age c1as$es.
Changes during and after EI Nino
It is obvious that certain population parameters
vary during normal upwellng periods. For example,
in M. donacium, differences in growth rate seem to
be related to temperature differences for 1980 and
1981. However, 1982-83 brought about very import-
ant changes in practically all parameters both at the
population and community level.
Unexpectedly, some of these changes occurred
several months before the Kelvin waves of warm
water hit Peru in the first half of October 1982 (Smith
(984). Similar observations of a premature impact of
EN were made by Tarazona et al. (in prep. b) in the
Bay of Ancon. Spawning of M. donacium which,
under normal conditions, occurs mainly from
December to March in Peru (Salgado and Ishiyama
1979) extended to the second half of 1982. As a
consequence, the main recruitment continued until
Oclober. Later, irregular recruitment occurred and
1987
the density of this species, which had dropped to a
very low value in June, once again increased until
January 1983. Nevertheless, the distribution of M.
donacium became more patchy. In February, when
EN reached its first peak of 26° C in Santa Maria,
nearly one-fifth of the specimens were found with
their valves open, and by March no survivors were
left in shallow water. Growth had stopped almost
completely by mid 1982, and there was no production
during the second half of that year. Apparently M.
donacium is sensitive to changes in temperature both
in positive and negative directions, reacting to lower
than normal temperatures with a decrease in growth,
and to higher than normal temperatures first with a
decrease, then with a halting of growth, and finally
with lotal mortality of the population.
Donax peruvianus revealed few year-to-year dif-
ferences in the population parameters before EN, but
clearly a slower growth in 1982-83 and hardly any
growth at all of the cohort recruited in 1983.
Mortality of Donax was not calculated, but many
empty valves among the Mesodesma shells in 1983
(Fig. 9b) indicated that few large individuals survived
the event. However, in contrast to M. donacium,
there was recruitment of D. peruvianus in 1983 which
enabled the population to survive.
Birkett and Cook (1987) show that the 1982-83
temperature anomaly caused irregular spawning by a
Donax serra population on the South African coast.
In this area, summer sea temperatures are usually
lowest because of strong upwelling, and spawning of
surf clams occurs in winter. However, temperatures
in summer 1983 climbed to 5°C above normal,
causing a significant proportion of the mature ani-
mals to spawn prematurely in summer, when they are
normally inactive, and delaying the winter spawning
maximum of 1983 for several months. The tempera-
ture anomaly affected principally the larval stages.
Settled spat occurred in very low numbers on the
beaches, resulting in a drastic recruitment failure in
1983. Significant numbers of spat reappeared only in
early summer 1984.
In contrast to the observations off South Africa, in
Peru by the end of 1983, as soon as four months after
the decrease in temperature, Donax peruvianus
density had climbed to the highest value observed
during the investigation. With only a relatively
modest figure, Donax became the first dominant.
When M. donacium disappeared from the Santa
Maria comm'unity, it was expected that one of the
species formerly ranked 2 or 3, or both of them,
would increase strongly and take the place of
Mesodesma so that some form of "neighbourhood
stability" (Gray 1977) would be maintained. In fact,
immediately after EN it seemed that the changes in
1987 Arntz et al.: Beach Community Structure Changes in Peru during El Nino 655
 The Benguela and Comparable Ecosystems
656 South African Journal of Marine Science 5
the two clam populations were going to shape the
further development of the community, whereas
other species appeared to remain insignificant. How-
ever, occasional sampling since EN (Tarazona, un-
published data) has revealed that there has not been a
single shift from the first to the second (or third)
dominant, though M. donacium has remained absent
from this community.
Not only in the months after EN did Donax never
reach densities similar to those of M. donacium
before the collapse of the latter species, but it has
remained at low densities up to the present. The same
observation has been made for E. analoga. Instead,
an impoverished community consisting mainly of
small opportunistic polychaetes became established,
and this has little similarity with the former Meso-
desma community. The start of this transitional
community of opportunistic species is shown in
Figure 4. Emerita also seems to do well during this
transitional stage. One conclusion which could be
drawn from this development is that the two clams,
despite their similar appearance and way of life,
never seriously competed for the resources at Santa
Maria beach. As D. peruvianus occurs in very high
densities in other sand communities, there may be
typical Donax and Mesodesma beaches, the differ-
ences of which remain yet to be identified. On the
other hand, Penchaszadeh (1971) mentions that,
during the period of Emerita dominance at Chilca
beach, there were no Mesodesma at all. In contrast,
during this study, Chilca was known among the
fishermen as one ofthe best places for collecting surf
clams. Therefore, the same beach has evidently been
populated by dense populations of mole crabs and
surf clams, although at different times. It seems likely
that Emerita are outcompeted by clams, or that the
clams modify the habitat in a manner that makes it
unsuitable for the mole crabs. There seem to be
long-term populations shifts which are not fully
understood at present.
Implications for fisheries and long-term stability
For the local fisheries, the events at Santa Maria
beach meant a change from a highly productive
community with easily accessible protein to a com-
pletely unproductive one. Changes of this type do not
appear to be unusual in the Peruvian-Chilean upwel-
ling system. Tarifeno (1980) mentions the case of a
Chilean beach now without any surf clams where
fishermen exploited a large commercial M. donacium
population about 30 years ago. It is possible that
former EN events caused massive population shifts
1987
along the coastline, as can be inferred from ancient
Mesodesma shell mounds in areas where the species
is absent nowadays (Fig. 9c). To the south, M. dona-
cium extended its area during the last EN as can be
seen from the landings in the surroundings of
Valdivia, Chile (400S) - SERNAP (1984). The large
Mesodesma populations present in the Department
of Lima until EN 1982-83 were quite recent, too;
colonization only started about 1975 (Tarazona,
unpublished data).
Before the last EN the northern limit of distribution
of M. donacium was near Trujillo (8° S), whereas it is
now south of l4°S. Donax peruvianus was affected
by the EN, but it increased in northern Chile (Soto
1985, Tomicic 1985). In the Pisco area, some other
clam species which were strongly affected by the EN
of 1982-83 have not recovered as yet (Arntz and
Valdivia 1985); among them are the bivalves Semele
spp. and Tagelus dombeii. Another notable, although
different, example is the scallop Argopecten purpu-
ratus, which increased tremendously during the last
EN near Pisco, Peru (l4°S) according to Wolff
(1984) and may fluctuate in a manner opposite to
that of M. donacium. Del Solar (1983) mentions
other examples amongst subtropical molluscan spe-
cies in northern Peru. At present it is not known
whether only exceptional Ninos such as that of 1982-
83 produce a shift of this nature, although this seems
very likely.
El Nino is the main factor affecting the medium-
and long-term stability of shallow-water communi-
ties off Peru and Chile, acting on a variety of spatial
and temporal scales. According to the strength (and,
possibly, frequency of occurrence) of EN, these
communities either show a high resilience, with a fast
recovery after each event, or they undergo decisive
changes which may last for decades. Even in the
latter case, the Peruvian and Chilean upwelling com-
munities within their total limits from the subtropics
to the subantarctic remain fairly persistent, although
their exact locality may fluctuate. The Humboldt
Current system with its countercurrents certainly
acts as an efficient transporter for meroplanktonic
larvae. Apparently, a certain flexibility will be re-
quired of the fishermen in the future too.
ACKNOWLEDGEMENTS
The authors thank C. Riemenschneider for re-
peatedly typing the manuscript, G. Dansauer, M.
PuIs and H. Westphal for the artwork, and three
referees for their helpful comments and corrections
1987 Arntz et al.: Beach Community Structure Changes in Peru during El Nino
of the English. Dr J. Alheit of the Alfred-Wegener-
Institut kindly presented the paper at the Benguela-
86 Symposium.
LITERA TURE CITED
ANSELL, A. D., McLUSKY, D. S., STIRLING, A. and
A. TREVALLION 1978 - Production and energy now in
the .macrobenthos of two sandy beaches in South West
India. Proc. R. Soc. Edinb. 768: 269-296.
ANSELL, A. D., SIVADAS, P., NARAYANAN, B. and
A. TRE~ ALLION 1972 - The ecology of two sandy
beaches In South West India. 3. Observations on the
population of Donax incamatus and D. spiculum. Mar.
BioI. 17(4): 318'-332.
ARNTZ, W. E. 1986· The two faces of EI Nino 1982-83.
Meere:.forschung 31( I): 1-46.
ARNTZ, W. E. and H. RUMOHR 1982 - An experimental
stu~y of macro benthic colonization and succession, and
the Importance of seasonal variation in temperate latitudes.
J. expl mar. BioI. £Col. 64{ I): 17-45.
ARNTZ, w. E. and H. RUMOHR 1986 - Fluctuations of
benthic macrofauna during succession and in an established
community. MeeresforsclllOlg 31: 97-114.
ARNTZ, W. E. and E. VALDIVIA 1985 - Incidencia del
fenomeno "EI Nino" sobre los mariscos en el litoral
peruano. In" EI Nino", su ImpaclO en la Fauna Marina.
Arnt?, W. [E.], Landa, A. and J. Tara7.0na (Eds). Boln
Inst. Mar, Peru-Callao, Spec. Issue: 91-101 (Proceedings
oj the EN Workshop within the Ninth Latin American
Conf<ress of Zoology. Arequipa. Peru, October 1983).
BIRKETT, D. A. and P. COOK 1987 - Effect of the Benguela
temperature anomaly, 1982-1983, on the breeding cycle of
Donax serra Roding. In The Benguela and Comparable
Ec~systems. Payne, A. I. L., Gulland, J. A. and K. H.
Bnnk (Eds). S. Afr. J. mar. Sci. 5: 191-196.
BREY, T. 1984· - Gemeinschaftsstrukturen, Abundanz, Biomasse
und Produktion des Makrozoobenthos sandiger Boden
der Kieler Bucht in 5-15 m Wassertiefe. Ber. Inst. Meeresk.
Kie/l2J: 124 pp.
BREY, T. 1986 - Estimation of annual PJ B-ratio and production
of marine benthic invertebrates from length-frequency
data. Ophelia. Suppl. 4: 45-54.
BREY, T. and D. Pi\ULY 1986 - Electronic length frequency
analYSIS. A revised and extended user's guide to ELEFAN
0, I, and 2. Ber. Inst. Meeresk. Kie/149: 76 pp.
CRISP, D. J. (Ed.) 1964 - The effects of the severe ice winter of
1962 63 on marine life in Britain. J. Anim. Ecol. 33(1):
165210.
CRISP, D. J. 1984 Energy now measurement. In Methodsfor
the SllIdy of Marine Benthos, 2nd Ed. Holme, N. A. and
A. D. Mcintyre (Eds). Oxford; Blackwell: 284-372 (/BP
Handbook 16).
DEFEO, O. 1983·-· Aspectos biocenologicos y de dinamica de la
poblacion de "almeja amarilla", Mesodesma mactroides
(Deshayes, 1854) en la zona de la Barra del Chuy, Depto.
de Rocha, Uruguay. M.Sc. thesis, Universidad de la
Republica, Montevideo: 94 pp.
DEFEO, O. 1?85a - Aspectos biocenol6gicos ydedinamicade
poblaclon de "almeja amarilla", Mesodesma mactroides
(Deshayes, 1854) en la zona de la Barra del Chuy, Depto.
de Rocha, Uruguay. I. Biocenologia. Con/rib. DeplO
Oceanogr. Univ. Repub. Montevideo 2(3): 50-75.
DEFEO, O. 1985b - Aspectos biocenologicos y de dimimica de la
657
poblacion de "almeja amarilla", Mesodesma mactroides
(Des hayes, 1854) en la zona de la Barra del Chuy, Depto.
de Rocha, Uruguay. 2. Dinamica de la poblaci6n. Contrib.
Depto Oceanogr. Univ. Repub. Montevideo 2(4): 76-98.
DEL SOLAR, E. M. 1983 - Variacion poblacionalde algunos de
los recursos vivos del mar por causa del fcn6meno de "EI
Nino" y el hombre. Boln Lima 27: 59-66.
G RA Y, J. 1977 - The stability of benthic ecosystems. Heigoiiinder
wiss. Meeresunters. 30: 427-444.
MOLLER, P. and R. ROSENBERG 1983 - Recruitment,
abundance and production of Mya arenaria and Cardium
edule in marine shallow waters, western Sweden. Ophelia
22(1): 33-55.
MOLLER, P., PIHL, L. and R. ROSENBERG 1985 - Benthic
faunal energy flow and biological interaction in some
shallow marine soft bottom habitats. Mar. Em/. Progr.
Ser. 27: 109-121.
OSORIO, C, BAHAMONDE, N. and M. T. LOPEZ 1967 - EI
limanche (Emerita analoga, Stimpson) en Chile. Boln
Mus. naco Hist.-nat. Chile 29: 61-116.
PAULY, D. and N. DAVID 1981 - ELEFAN I, a BASIC
program for the objective extraction of growth parameters
from length frequency data. Meere~forschung 28(4):
205-211.
PAULY, D. and G. GASCHOTZ 1979 - A simple method for
fitting oscillating length growth data, with a program for
pocket calculators. ICES eM. 1979/ G:24: 26 pp. (mimeo).
PENCHASZAUEH, P. E. 1971 - Observaciones cuantitativas
preliminares en playas arenosas de la costa central del Peru
con especial referencia a las poblaciones de muy-muy
(Emerita analoga) -- (Crustacea, Anomura Hippidae).
Doe. In! Oflc. Ciene. UNESCO Am. Lat., Montevideo 6:
16 pp.
'RACHOR, E. and S. A. GERLACH 1971l - Changes of
macrobenthos in a sublittoral sand area of the German
Bight, /967 to /975. Rapp. P.-v. Reun. Cons. perm. into
Explor. Mer 172: 41ll·431.
ROSENBERG, R., ARNTZ, W. E., CHUMAN DE FLORES,
E., FLORES, L. A., CARBAJAL, G., FINGER, I, and
J. TARAZONA 19l13 - Benthos biomass and oxygen
deficiency in the upwelling system off Peru. J. mar. Res.
41(2): 263-279.
ROSENBERG, R., LARSON, U. and L. EDLER 1986 --
Eu~rophication in marine waters surrounding Sweden -. a
revIew. Rep. noln. Swed. En vir. Prot. Bd 3054: 137 pp.
RUMOHR, H. and W. E. ARNTZ 1982 - The "Benthosgarten"
- a new approach for the study of soft bottom com-
munities. Meeresforschung 29(4): 225-238.
SALGADO, I. and V. ISHIYAMA C 1979 - Cicio de madurez
sexual y desove de la macha Mesodesma donacium. Revta
Fac. Cienc. Univ. naco Lima 71( I): 20-28.
SERNAP (Servicio Nacional de Pesca, Chile) 1984 - Anuario
estadistico de pesca 1984. Ministerio Economia, Fomento
y Reconstruccion, Santiago-Chile: 82 pp.
SMITH, R. L. 1984 - Coastal currents off Peru during El Nino of
1982-83: prenatal to senescent conditions. Trop. Ocean-
Atmos. Newsl. 28: 7-8.
SOTO, R. 1985 - Efectos del fenomeno El Nino 1982-83 en
ecosistemas de la 1 Region. Investigacilm pesq., Santiago
32: 199-206.
TARAZONA, J., ARNTZ, W. E., CANAHUIRE, E.,
la AYALA, Z. and A. ROBLES 1985 - Modificaciones
producidas durante "EI Nino" en la infauna bent6nica de
areas someras del ecosistema de anoramiento peruano. In
"EI Ni;;o': su Impactoen la Fauna Marina. Arntz, W. [E.],
Landa, A. and J. Tarazona (Eds). Boln Inst. Mar. Peru-
Callao, Spec. Issue: 55-63 (Proceedings of/he Ninth Latin
 The Benguela and Comparable Ecosystems
658 South African Journal of Marine Science 5 1987

American Congress of Zoology. Arequipa. Peru. October
1983).
TARAZONA, J., SALZWEDEL, H. and W. [E.] ARNTZ (in
preparation a) - Positive effects of"EI Nino" on macro-
zoobenthos inhabiting hypoxic areas of the Peruvian up-
welling system.
TARAZONA, J., SALZWEDEL, H., ARNTZ, W. [E.] and
L. ROMERO (in preparation b) - Oscilaciones en el
macrobentos de aguas someras de la costa central del Peru
inducidas por "EI Nino" 1982-83. In Proceedings of the
Chapman Conference on EI Nino. Guayaquil/ Ecuador.
October 1986.
TARIFENO, S. E. 1980 - Studies on the biology of the surf clam
Mesodesma donacium (Lamarck, 1818) (Bivalvia: Meso-
desmatidae) from Chilean sandy beaches. Ph. D disserta-
tion, University of California, Los Angeles: 229 pp.
TOMICIC, J. J. 1985 - Efectosdel fen6meno EI Nino 1982/83 en
las comunidades litorales de la Peninsula de Mejillones.
Investigaci6n pesq., Santiago 32: 209-213.
WEIGELT, M. and H. RUMOHR 1986 - Effects of wide-range
oxygen depletion on benthic fauna and demersal fish in
Kiel Bay, 1981-1983. Meeresforschung 31: 124-136.
WOLFF, M. 1984 - Impact of the 1982-83 E1 Nino on the
Peruvian scallop Argopecten purpura/Us. Trop. Ocean-
Atmos. Newsl. 28: 8-9.
ZIEGELMEIER, E. 1964 - Einwirkungen des kalten Winters
1962/63 auf das Makrobenthos im OSlleil der Deutschen
Buehl. Helgoltinder wiss. Meeresunters. 10: 276-282.
ZIEGELMEIER, E. /970 - Ober Massenvorkommen ver-
schiedener makrobenthaler Wirbelloser wah rend der
Wiederbesiedlungsphase nach Schadigungen durch "kata-
strophale" Umwelteinfliisse. Helgoliinder wiss. Meeres-
unters. 21: 9-20.