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Growth Increment and Stable Isotope Analysis of Marine

Bivalves: Implications for the Geoarchaeological
- -'
Record of El Nio

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Harold B. Rollins
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Department of Geology and Planetary Science, University of Pittsburgh, Pittsburgh, PA 15260

Departrrumt of Anthropology, Cornell Uniuersity, lthaea, NY 14853
Uwe Brand
Department ofGeology, Brock Uniuersity, Sto Catharines, Ontario, Canada
Juditb C. Rollins
329 Bellwalt Dr., Bridgeville, PA 15017
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The 1982-1983 El Nio event afforded the opportunity to develop eriteria for the recognition ofancisnt Ei
Nios using mollusks from archaeological sites along coastal South America. A combination of growth
increment and stable isotope analyses indicated that elevated sea surface temperatures during large seale
El Nios leave a record decodable from the growth patterns of selected bivalve shells. The intertidal
venerid Chione subrugosa displayed a pronounced break in the valve margin profile following the 1982
1983 vent but provided an inconsistent stable isotope pattern. The subtidal carditid Trachycardium
procerum, on the other hand, preserved a discernible and diagnostic growth interruption as weU as an
expected trend in stable isotope indicators of salinity and temperature change.
We conclude that some of the major culturally disruptive El Nio events can be recognized in the
geoarchaeological record by these techniques, especially ifandllary information. such as faunal distribu
tion pattems, are also considered. Perhaps the most serious constraint upon application ofthis
involves microstratigraphic resolution of shell midden deposits. Stratigraphic sampling ofmidden mate
rial should be accompanied, ifpossible, by sampling ofproximal natural strata. The chances ofdiscovery of
major El Nio perturbations in the geoarchaeological record of shell middens is enhanced by the cata
strophic nature of such events and by the indication that major El Nios have a high probability ofbeing
closely spaeed in time.
The predominantly maritime-based econo
mies of coastal Peru have existed for thou
sands of years (Moseley, 1975; Sandweiss et
al., 1983; Richardson, 1981). For at least the
last 5000 years El Nio must have been, as it
is now, a major infl uence upon coastal occupa
tion of that area (Rollins et al., 1986a). The
recognition of ancient large-scale El Nio
events would certainly aid our understanding
of the ecological vagaries that must have, in
part, affected the growth and development of
these coastal economies.
The 1982 1983 El Nio event has afforded
us the opportunity to develop criteria for the
recognition ofancient large-scale El Nio per
turbations, utilizing the widespread and
abundant shell midden deposits along coastal
Peru. This paper discusses the application of
growth increment and stable isotope analyses
of bivalve shells as a tool for recognition of
ancient El Nio events.
TBE 1982-1983 EL NIO
The 1982-1983 El Nio event was proba
bly the most devastating Pacific Ocean cli
matic perturbation of this century (Arntz,
1986). Historical records of El Nio events
extend back to the 1700s and, using any avail
able index, the 1982-1983 El Nio ranks as
Geoarchaeology: An International Journal, Vol. 2, No. 3, 181-197 (1987)
CJ1987 by John Wiley & Sons, Ine. eee 0883-6353/87/030181-17$04.00
one of the worst (Cane, 1983; Quinn et aL,
1978). The event was atypical in many re
spects. Thermal anomalies (as measured by
SST-sea surface temperature) were extraor
dinarily high and the effects ofthe event did
not hit coastal South America untillate sum
mer, when local SST would normally be de
clining to its lowest value. Caviedes (1984)
presented a detailed chronology of the 1982
1983 events that affected the coast of South
America. In June of 1982, the coast of central
Chile received high rainfall with repeated
passage of South Pacific winter depressions.
By October, 1982, excessive rainfall occurred
in coastal Ecuador and southern Colombia
but did not affect the region south of about
latitude 2 degrees south. However, September
and October were times of dramatic rise in
SST along the South American coast. The tor
rential rainfall moved southward during De
cember, following the abnormal southern dis
placement of the intertropical convergence
zone (lTCZ). Premature andexcessive rainfall
reached northern Peru by J anuary, causing
extensive flooding and landsliding. While
there was rainfall in the desert of northwest
Peru, drought struck the agricultural regions
of southern Peru (Rasmussen and Wallace,
1983). Violent storms continued along the
coasts ofEcuador and norther Peru until mid
June. Sorne degree of seasonal atmospheric
normality was restored by the end of June,
1983 (Caviedes, 1984). Although the ITCZ
had passed beyond northern Peru, air temper
atures were still higher than usual and the
SST of coastal waters north of Callao, Peru
(about latitude 12 degrees south; Figure 1)
was as much as 8C aboye normal. The Paita
coastal station (about latitude 5 degrees
south) showed signs of recovery of SST nor
mality by July, 1983 when the temperature
decreased to 20C (Barber and Chavez, 1983).
Ecological disruption following the onset of
the 1 9 8 2 ~ 1983 El Nio was dramatic but not
at all unexpected. Modification of the normal
upwelling of cold nutrient-rich water adjacent
to coastal South America inevitably affects
populations of phytoplankton, fish, and sea
birds. The response of larger marine pelagic
organisms to SST anomalies can be very
rapid, ostensibly reflecting a behavorial adap
tation triggered by temperature change,
when there is actually a complex linkage
among nutrient supply, light intensity, tem
perature, productivity, and food (Barber and
Chavez, 1983). The stresses created by
reduced food availability are apparentIy am
plified up the food web, leading to decreased
growth rates and reproductive failure most
noticeable at higher trophic levels. These cat
astrophic ecological disturbances are often
unappreciated over short spans oftime due to
the spatial displacement of many species fol
lowing the onset ofEI Nio. Local "catches" of
sorne fish and shellfish may actually increase
due to movement by currents ofdisplaced pop
ulations into shallower, more accessible
coastal regions or due to the temporary intro
duction of thermally anomalous "exotic" spe
cies. Arntz (1986) noted, for example, that the
1982-1983 El Nio caused rapid growth and
high den sities of scallops and sorne other in
vertebrate species. On the other hand, nter
tdal and shallow subtdal mussel banks and
kelp forests were destroyed by the evento Hu
man inhabitants of these coastal areas who
are dependent upon gathering of nearshore
invertebrates for food were hardest hit by the
1982-1983 catastrophe. This most likely has
been true in this region for every large-scale
El Nio event over thousands of years.
The economically important bivalve species
of coastal Peru are, for the most part, sessile
benthic and thus incapable of short-term mi
gration with changes of water currents and
temperature. Over the long term, however,
such species exhibit ecological resilience due
to their"high reproductive recruitment poten
tial. Although scattered anecdotal informa
tion is available concerning the effects of El
Nio upon shellfish harvests, Httle research
effort has been directed toward understand
ing physiological and morphological re
sponses of various shellfish species to the eco
logical stresses ofEI Nio. Sessile bivalve spe
ces are sensitive indicators of changing
coastal condi tions and record, on a daily
andlor subdaily basis, a chronology of envi
ronmental perturbations. This record is mani
fested in the pattern of shell growth incre
182 VOL. 2, NO. 3
- - - - ~ - - - - - - - - - - - - - - - - - - ~ ~ ~

o 100 200
Figure l. Map ofPeru showing cities mentioned in texto
ments, types of shell microstructure, and
profile of the value anterior margino
The bivalve specimens utilized in this study
were harvested alive or purchased from ven
dors at several sites along coastal Peru, dur
ing a series of collecting trips in 1984. CoBect
ing stations ranged from Tumbes, in extreme
northwestern Peru, tri the Paracas pennsula,
about latitude 15 degrees south (Rollins et al.,
1986b; Figure 1).
Following preliminary screening of several
harvested bivalve species, 35 specimens ofthe
intertidal Chione subrugosa and 15 speci
mens of the subtitdal Trachycardium pro-
cerum were selected for detailed growth incre
ment analysis. Description of the techniques
of bivalve growth increment analysis are
readily available from the geoarchaeological
literatur and will only be briefly described
here (Deith, 1983; Koike, 1980; Rollins et al.,
in press). Individual valves were cut with a
Felker rock saw along the axis of maximum
growth (normal to surface growth lines).
Fragile valves were embedded in Epon 815
epoxy resin prior to cutting (Kennish et al.,
1980). Cut valves were ground and polished
using a Buehler Isomet lapidary machine and
then etched in 5% HCl for approximately one
minute. Etching time varied according to
species and condition of valves. The etched
valves were carefully washed, air-dried, im
mersed in acetone and quickly placed upon
pieces of sheet acetate (0.1 mm thick). After
drying, the acetate peel replicas were
mounted between glass microscope sUdes. Ac
etate peel replicas were studied and photo
graphed using a MPV-2 microscope and auto
matic camera. Growth increment counts were
made upon assembled photo mosaics.
Stable isotope analysis of bivalve shells is
also widely used in geoarchaeological studies,
mainly as a technique for ascertaining season
ofharvesting (Shackleton, 1983; Bailey et al.,
1983; Jones et al., 1984; Deth, 1986; reviewed
by Rollins et al., in press). Killingley and
Berger (1979) were able to detect upwelling
events by stable isotope analysis ofCalifornia
mussel shells. We know of no other study,
however, which has attempted to use stable
isotopes ofmolluscan shells as indicators ofEI
Nio sea surface temperature perturbations.
Prior to stable isotope analysis all bivalve
shells were initilly cleaned of adhering or
ganic and inorganic material. Then the shells
were cut along the axis ofmaximum growth in
the manner described aboye and subjected to
further cleaning by immersion in an ultra
sonic bath of de-ionized water (to minimize
contamination) for 15 minutes. Additional
cleaning consisted of brief immersion in 15%
HCI, rinsing with de-ionized water, and sub
sequent air-drying (Brand and Veizer, 1980).
Shell samples were taken (by drilling and
sawing) at specific intervals along the axis of
maximum growth in order to traverse the
growth before, during, and after the 1982
1983 El Nio event (as indicated by examina
tion ofvalve margin pro file and growth incre
ment pattern). These samples were powdered
and dissolved in 5% HCI, in order to minimize
the leaching of elements from material that
was not removed during the cleaning process.
All samples were analyzed for Ca, Mg, Sr,
Na, Mn, Fe, Al, Ni, Cu, and Zn usng an auto
mated and computerized Varan 1475 atomc
absorption spectrophotometer (see Brand and
Veizer, 1980; Brand and Hinsperger, 1986 for
complete details of sample preparation and
analysis). Trace element compositions were
calculated on a 100% carbonate free basis to
minimize trends due to fluctuations in
organic matter content ofthe shells. Accuracy
of chemical values was determined by analyz
ing N.B.S. standard rocks (633,634,636) and
precision of data was calculated using dupli
cate analyses.
The shell samples were also analyzed for
oxygen and carbon stable isotopes. Powders
were reacted with 100% phosphoric acid at
50C for 10 minutes, and evolved gas was
analyzed on a V.G. 903 Micromass spectrome
ter. The isotope ratios are expressed in 8 rela
tive to PDB and corrected for oxygen-17.
Chemical data were evaluated using the
statistical packages (e.g., SPSS) on a Bur
rough's B7900 computer, and with the Stat
view program on an Apple Macintosh com
putero Graphs were computer-generated.
A total of 14 samples from three specimens
ofChione subrugosa and 19 samples from four
specimens of Trachycardium procerum were
analyzed for Ca, Mg, Sr, Mn, Na, Fe, Zn, Cu,
and Ni. AIso, a representative suite of15 sam
pIes was analyzed for 8
0 and 8
Growtb Increment Analysis
Chione subrugosa and Trachycardium pro
cerum display different patterns of growth.
The former has a reflected mantle margin, as
do other venerid bivalves, and easily counted
arcuate growth increments are deposited
(Figure 2). Trachycardium procerum pos
sesses a nonreflected mantle margin and
growth increments intersect the shell margin
at high angles (Figure 3). In Chione subru
gosa daily growth increments were prominent
while semi-daily growth increments were
most distinct in Trachycardium procerum.
Detailed analysis of the daily and semi-daily
growth increments of bivalve shells has
proven to be a useful indicator of short and
long term environmental stress, intensity of
seasonal zonation, and habitat relative to
onshore/offshore depth gradients (Richardson
et al., 1980; 1981; Barker, 1964; Jones, 1983;
Rhoads and Lutz, 1980; Hall et al., 1974; Ev
ans, 1975; Clark, 1975; Kennish and Olsson,
1975; Thompson, 1975; Pannella and Mac-
VOL 2, NO. 3
--...0.1 mm
Figure 2. A-C. Chione subrugosa (photomicrographs of acetate peels). A. Specimen
# Ptl-10, collected 4-16-84, Tumbes, Peru. Note distinct daily lines and subtle,
dscontinuous sub-daily lines. Crossed-Iamellar microstructure (X) is in typical un
stressed pre El-Nio orientation. Growth margin of the shelI is to the left. B. Same
specimen. Area immediately below El Nio break. Note transgressive crossed
lameIlar microstructure (X) invading upper composite prismatic layer. C. Specimen #
FPr-1, collected from shell midden, Pampas las Salinas, Peru. N arrow break in valve
profile is typical of a storm break, not a major El Nio disturbance. Recovery was rapid
following the storm break. Growth margin of the shell is to the right. D-E.
Trachycardium procerum. Arrows indicate onset of El Nio break. D. Specimen #
TPl-6, collected 4-18-84, ChicIayo, Peru. E. Specimen # 2TP2-1O, collected 12-1-84,
Trujillo, Peru.
Figure 3. Trachycardium procerum (Speciman 1/: TP3-l), collected 4-20-84,
Chimbote, Peru, A-B. Acetate peel, negative print (scale 1 mm). Note broad
shallow El Nio break in valve profile and increase in crossed-Iamellar microstruc
ture at the val ve margino Anterior tip is to the right in all of the photographs.
Semi-daily growth incrementa are shallowly inclined below the El Nio break. Ar
rows indicate 307 days (614 increments). C. Photomicrograph (scale = 0.1 mm).
Enlargement of area C in Figure 2A. Note mass'Ve crossed-lameIlar microstructure
and nearly horizontal growth increments. D. Photomicrograph (scale == 0.1 mm) of
area D in Figure 2A. Cross-lameIlar microstructure still present, but growth incre
ments have resumed normal inclination to the valve margino
Clintock, 1968; Ekaratne and Crisp, 1982). colored bands, the result of calcium carbonate
There is general agreement that bivalve deposition when val ves are open and the for
shells are gauges of a hierarchy oflunar and mation of a thin dark line, presumably due to
solar rhythms, as well as extrinsic environ anaerobic dissolution of shell material, when
mental events. An individual growth incre valves are c10sed (Lutz and Rhoads, 1977).
ment consists of a couplet of light and dark Intertidal bivalves (e.g., Chione subrugosa)
186 VOL 2, NO. 3
therefore have incremental patterns that re
flect both lunar tidal (exposure and immer
sion) and solar day rhythms where valve cIo
sure accompanies exposure and increased
input ofheat Oight). Sorne controversy exists
over the temporal significance of single
growth increments in bivalve shells. Richard
son et al. (1980, 1981) have'made a strong case
for tidal exposure and temperature control
ling the expression of growth increments.
Prominent growth lines are the result oftidal
exposures coinciding with elevated tempera
tures, as when low tide occurs durir.g daylight
hours. Alternating strong and weak lines oc
cur when one low tide is in early morning and
the other is in mid-afternoon (Richardson et
al., 1981). We suspectthatsuchfactorslead to
a pronounced difference in strength of growth
lines so that bivalves which are totally ex
posed at low tide will appear to form one line
per day whenever one low tide coincides with
daylight hours. Counting of growth lines is
difficult and the tendency is to observe and
record only the more prominent lines. Pre
sumably, growth line counts on high nter
tidal bivalves will tend to emphasize daily
perodicities whereas counts on lower inter
tidal and subtidal bivalves stress semi-daily
intervals. This may explain why daily lines
have been commonly recognized in venerids
such -as Mercenaria mercenaria, Meretrix
lusora, and Chione spp., which inhabit inter
tidal mud flats or lagoons (Koike, 1973;
Pannella and MacClintock, 1968). The sub
tidal carditids, in contrast, tend to display
nearly equal development of semi-daily lines,
corresponding to each low tide. Our analyses
of Trachycardium procerum and Chione sub
rugosa support this interpretation.
As noted by many workers (e.g., Thompson,
1975), shell increments may al so be grouped
into alternating close and wide-spaced bun
dles that correlate with semi-monthly neap
and spring tides. The tidal patterns are gener
ally less obvious in subtidal bivalves (e.g.,
Trachycardium procerum). In more temper
ate regions annual patterns of increment de
position reflecting seasonality may also be
Specific environmental or biological
stresses may also result in pronounced growth
breaks in bivalve shells. Such stresses may
accompany storms, temperature shocks (hot
or cold), or spawnng. Breaks due to hot and
cold shock may be quite similar, exhibiting
sudden bunching of growth increments at the
onset, with slow recovery. Storm breaks, how
ever, typically display rapid onset and recov
ery, and may result in the depositional incor
poration of sediment grains at the base of a
narrow notch in the shell margino Excellent
discussions of the types of growth patterns
and breaks are provided by Kennish and
Olsson (1975) and Cunliffe (1974). Kennish
and Olsson (1975) determined that the major
controlling factors affecting the growth ofthe
venerid clam Mercenaria mercenaria were
temperature, tides, substrate type, water
depth, and age ofthe individual. Environmen
tal stress also may be indicated by irregulari
ties in the shell microstructure and in the
profile of the valve margino Kennish and
Olsson (1975) noted that thermal shock
breaks in venerids may cause a transgression
of massive crossed lamellar microstructure
into the outer prismatic or composite pris
matic microstructure. Wide notches may oc
cur on the valve margins as a result of the
temporary withdrawal of the mantle during
prolonged stress. They are quite distinct from
the narrow storm-induced notches mentioned
aboye and may result from long-term clima tic
perturbations such as El Nio events. Under
such conditions the ventral margin of the
valve may be blunted with a precipitous slope
(Rollins et aL, 1986b).
Conspicuous El Nio-induced growth
breaks were present in all the specimens of
Chione subrugosa collected in 1984 from
Tumbes (Figure 4). The growth breaks dis
played rapid onset and slower, often erratic,
recovery. Most specimens did not achieve to
tal recovery and the ventral shell margins are
strongly blunted. In contrast, the ventral
margins of unstressed specimens of Chione
subrugosa are much thinner, as indicated by
comparison of these specimens with a collec
tion of 194 Chione subrugosa valves from an
archaeological site at Valdivia, Ecuador.
Sixty-one percent of the Ecuadorian shells

: : r = ~
Figure 4. Chione subrugosa. Hachure marks indcate position ofEI Nio breaks.
A-B. Specimen # PTl-8, Tumbes, Peru, collected 4-16-84. C-D. Specimen #
Vald-2, Valdivia midden, Cut J, coastal Ecuador. E-F. Specimen # PTl-17,
Tumbes. Peru, collected 4-16-84. G-H. Specimen # PTl-19, Tumbes, Peru, col
lected 4-16-84. I-J. Specimen # PT1-10, Tumbes, Peru, collected 4-16-84. K-L.
Specimen # 2PTl-6, Tumbes, Peru, coUected 11-25-84. M-N. Specimen #
2PTl-ll, Tumbes, Peru, collected 11-25-84.
had smooth profiles, with no stress breaks
(Figure 4C, D). When breaks were observed
they were not the major interruptions de
scribed above (Rollins et al., 1986b).
Moreover, the Ecuadorian shells that had
stress breaks near the ventral margins of the
shells showed rapid recovery and a thin ven
tral margin was reestablished after the
growth interruption in the val ve profile. The
El Nio-stressed specimens did not exhibit
such recovery.
Growth increment counts taken on the
photomosaics of acetate peel replicas of
Chione subrugosa extended from the ventral
tips to the beginning of the major growth in
terruption. Increment counts ranged from
VOL 2, NO. 3
300-375 (average of 328) for specimens col
lected alive on April 16, 1984. The increment
counts on another specimen of Chione sub
rugosa killed on November 25, 1984 totaled
531 (Table 1). Counting backward from the
ventral margin and assuming one increment
addition per day places the time of onset of
major stress on C hione subrugosa in late May,
1983. This is coincident with the maximum
SST anomalies measured at the Peruvian
ports of Paita and Chicama when water tem
peratures reached 10C aboye normal. Such
increment counts are probably underesti
mates beca use ofthe likely anaerobic dissolu
tion or nondeposition of increments during
intervals of maximum stress. Our results
closely agree with the growth increment
counts of Pallant (in press), who used growth
breaks and the absence of sub-daily lines to
infer stressful habitats of Chione subrugosa
in a study of environmental changes in Costa
Rica. Our data support Pallant's interpreta
tion as we noted that sub-daily lines were
deposited only prior to the onset ofthe El Nio
SST anomalies. Stress resulted in closely
spaced lines of about equal strength.
Specimens of Chone subrugosa exhibited
slow and erratic recovery from the El Nio
induced stress (Figure 5). The growth nter
ruptions recorded by notches in the valve
profiles, unlike storm breaks, sometimes
spanned several months and presumably in
dicate extended periods of partial mantle re
tracton. The trauma of the El Nio-induced
stress is further indicated by a lack of corre
spondence, among individuals, in the amount
of post-break shell deposition. In fact, many
specmens displayed as much post-stress
growth directed inward as there was horizon
tally along the radial growth gradient. This is
indicated by the scatter and overlap of the
meaSUTements plotted in Figure 6. The er
ratic pattern of growth recovery in this spe
cies substantiates the interpretation of a
stress-induced break in contrast to sorne type
of ontogenetically programmed slow-down of
shell growth.
Increment analysis of Trachycardium pro
cerum corroborates the results of the Chione
subrugosa counts. Comparable counts from
the ventral tip to the initiation of the growth
break were about double those in Chione
subrugosa (Table 1), and denoted major stress
commencing on April 30,1983 and June 20,
1983 for the two specimens examined. These
dates are well within the interval of elevated
El Nio-induced stress profiles are less ac
centuated on Trachycardium procerum than
on Chione subrugosa and may be subtle
bunching of growth lines or broad shallow
depressions (Figure 3). Recovery of Trachy
cardium procerum following El Nio stress
involved resumption of relatively normal
growth and the addition of several centime
ters of shell material. Only a few specimens
displayed pronounced inward growth but in
these cases the slopes ofthe valve profiles are
more gentle, presumably due to the nonre
flected mantle margin in this species. During
major stress the angle of inclination of the
growth increments to the valve surface is only
about 10 degrees. Normally (i.e., under
nonstressful conditons) the inclination angle
is about 35 degrees.
Trachycardium procerum showed trans-
Table l. Counts of Growth Increments-Ventral Margin to Onset
of El Nio Break.
Chione subrugosa
Date of # of # of Onset of El Nio
Specimen # Locality Death Increments Days Break-Date
2PI'1-11 Tumbes 11/25/84 531 531 6/16183
PI'1-19 Tumbes 4/16/84 325 325 5/28/83
PI'1-1O Tumbes 4/16/84 300 300 6/22/83
PI'1-17 Tumbes 4/16/84 375 375 4/8/83
Trachycardium procerum
Chiclayo 4/18/84
Chimbote 4/20/84
Figure 5. Chione subrugosa. Specimen # PTl-19, collected 4-16-84, Tumbes, Peru.
A. Acetate peel, negative print (scale = 1.0 mm). Arrow indicates onset ofEI Nio
break at 325 days (325 increments) prior to death. Note extreme bunching of growth
increments from the initial break in valve profile to the anterior tip ofthe valve. B-D.
Photomicrographs (scale = 0.1 mm). Enlargements of the stress profile shown in
Figure 4A.
gressive development of crossed lamellar support to the recent studies of Cerastoderm:;
shell microstructure as far as the shell margin edule, another carditid clam (Richardson et
in association with the El Nio break. In al., 1980, 1981). Inthat species, tidallevel was
Chione subrugosa the inner crossed lamellar determined to be a major factor influencing
layer also extended part way into the outer the rate of growth and more rapid growth
composite prismatic layer concomtant with correlated with longer periods of immersion.
the El Nio break, but never carne close to the
shell margin (Figure 2).
Stable Isotope Analysis
The nearly equal strength of semi-daily
growth increments in Trachycardium pro For the most part, the elemental analysis of
cerum follows the pattern expected for low the shells of Chione sub rugosa and Trachy
intertidal and subtidal bivalves and lends cardium procerum resulted in anomalous or
190 VOl. 2, NO. 3

E x
> 1

\ NOV 25, '84


\ y= 0476X + 347


r = 04

NOVEMBER 25, 1984 (N= 15)
X APRIL 16, 1984 ( N-20)
le X
16, '84
30 31 32 33 34 35 36 31 38 39 40 41 42 43
TH (mm)
Figure 6. Two collections ofChione subrugosa. Graph shows lack of correlation between amount of post-El
Nio shell growth (as measured by DVM-horizontal distance from El Nio break to growing margin ofthe
shell) and total shell height (measured from umbo to growing margin of the sheJl). The El Nio-induced
shock resulted in long-term growth interruption. DVM does not record the amount of inward-directed, or
vertical, component of growth, however.
insignificant chemical differences between
the pre- and post-El Nio events. However,
SrlNa and stable isotope trends in the shell
composition ofTrachycardium procerum were
indicative of changing oceanographic condi
tions during the El Nio evento In contrast,
Chione subrugosa exhibited a statistically in
significant SrlN a and stable isotope pattern
(compare Figures 7 -12 and Table ID.
Upwelling currents are generally enriched
in the light carbon isotope beca use the total
dissolved inorganic carbon contains more
C-12 than C-13. Therefore, the incursion of
warm surface waters ofthe equatorial counter
current into the colder waters ofthe Peruvian
Province should result in heavier values of
C in the carbonate shell material precipi
tated during the incursion (Killingley and
Berger, 1979). At the same time, the warmer
waters should result in lighter 8
0 values in
molluscan shell carbonate.
The anomalous values of both 8
C and
0 exhibited by specimens of Chione
subrugosa (Figures 8, 9) are consistent with
the habitat ofthis species. Lagoonal mud flats
are subject to frequent episodic localized tem
perature spikes and we anticipate that the
shell record of such an intertidal species
would exhibit a virtually indecipherable pat
tern of temperature sensitive stable isotope
In contrast, the elemental and isotopic com
oe PTI-17
6. PT'-8
Relative Growth Increment
Figure 7. Sr/Na-growth increment distribution dia
gram for three specimens of Chione subrugosa. Solid
symbols represent pre-EI Nio shell growth. EN is the
onset of the El Nio-induced shock, as indicated by
growth abnormality along the valve margin profile.
Open symbols represent shell carbonate deposition dur
ing and after the El Nio shock.
Table 11. Unpaired student t-Test of Sr/Na (wt) in
Chione subrugosa and Trachycardium procerum
growth increment samples.
Sr/Na (wt) X
Allochem N EL t-value
C. subrugosa 0.234 0.248 0.756 ,,;;; 0.375
T. 0.413 0.271 -3.796 ,,;;; 0.005
N '= pre-El Nio growth increment samples; EL '= post
and El Nio increment shell samples.
positions of Trachycardium procerum appear
to reflect the physiochemical conditions of the
ambient seawater. The Sr/Na ratios in the
shell incrcment samples decrease after the El
Nio event (Figure 10). The pre- and post-El
Nio values are significantly different at the
99.5% confidence level (TabIe II). Similar
trends are also depicted by the 8
C composi
tions of Trachycardium procerum, and the
heavier 8
e values correlate with the ele
vated SST along coastal Peru (Figure 11).
Concurrently, the 8
0 values are lighter in
the post-El Nio precipitated shell carbonate
(Figure 12), presumabIy an indication of the
warmer counter current water.
+ 0.5
-1. 5
Relative Growth Increment
Figure 8. o
C-growth increment distribution dia
gram of C hione subrugosa. Explanations and symbols as
in Figure 7.
Water salinities and temperatures can be
calculated with:
Salinity (1.0, ppt) = -5.037 In A + 28.627,
where A is the Sr/Na (wt.) ofindividual arago
nite samples (Brand, 1984) and with:
Toe = 19.0 -3.52(8
+ 0.14(8
- 8
where the 8
is the
0 ratio of the mol
luscan aragonite and 8
is the 160p80 ratio
of the ambient seawater (Grossman and Ku,
1981). Application of these equations to the
Sr/Na values of Trachycardium procerum
(Figure 10) resulted in calculation of an aver
age pre-El Nio water salinity of about 33 ppt
and a post-El Nio average value of 35 ppt.
Similarly, the calculated water temperature
VOL 2, NO. 3
1.4 ..-----------...,
1.6 \
\ a
rr l
Relative Growth Increment
Figure 9. &lSO-growth increment distribution di a
gram ofChoine subrugosa. Explanations and symbols as
in Figure 1.
increased from a preEI Nio average of 17C
to 22C after the incursion of the warm equa
torial counter current.
We believe that the results of this study
have the potential for establishing a baseline
for recognition of ancient El Nio events, uti
lizing the abundant and widespread shell
midden material along coastal South Amer
ica. For the most part, species that are har
vested today along coastal South America are
available from middens and associated natu
ral deposits that reach back through thou
sands of years of maritime culture, although
not without occasional widespread changes in
species distribution patterns (Rollins et al,
Recognition of large scale El Nio events
from analysis of shell midden material will
have to proceed with caution, however. We
suspect that a detailed chronology of El Nio
will never be attained by these techniques,
but we are optimistic that sorne of the major
culturally disruptive El Nio events can be
. recognized in the geoarchaeological record.
As such, we urge that attention be paid to
the following assumptions and constraints:
1. Clearly, detailed growth increment and
stable isotope analyses of bivalve shells are
limited by time and cost constraints. The type
of growth increment analysis employed in
this studyutilizing numerous photomosaics of
acetate peel replicas is labor intensive and
any study must necessarily be restricted to a
relatively small number ofbivalve shells. By
the same token, geochemical analysis of bi
valve shells is constrained primarily by cost
and the large number of samples required per
Techniques for more rapid processing of
molluscan shell material will be essential.
The results of our study suggest that the pro
file of the valve margin will provide a quick
and relatively easy method for recognition of
El Nio-induced stress among large collec
tions of bivalve sheIls. The El Nio "signa
tures" recognized in Chione subrugosa and
Trachycardium procerum involved unique
patterns of growth disruption that could be
c1early separated from shorter-term environ
mental stress, such as isolated storm events.
The use of the valve profile technique is en
hanced by shell growth in tropical and sub
tropical latitudes where strong seasonal
growth disruptions are absent.
We suspect that the role of detailed growth
increment and stable isotope analyses will be
restricted to occasional checks of randomly
selected shells from midden material (and as
sociated natural deposits) and confirmation of
suspected El Nio-induced patterns.
2. Our study demonstrated a strong habi
tat andJor biological constraint upon the use
of geochemical analysis of bivalve shells for
recognition ofEI Nio-induced stress. Wecan
not expect that intertidal species will respond,
in terms of stable isotope ratios, similarly to
subtidal species. The results ofour study sug
gest that subtidal species will be more reliable
geochemical prospects, but even intertidal
species can be used if detailed growth incre



o. TP3 -1
c.. 2TP2 -10
Q. TPI -6
o. TPI -9
Relafive Growfh Incremenf
Figure 10. Sr/Na-growth increment distribution diagram
for four specimens of Trachycardium procerum. Explanation
and symbols as in Figure 7.
tI. 2 ....-------------...,
Relaflve Growth Increment
Figure 11. &13C-growth increment distribution dia
gram of Trachycardium procerum. Explanations and
symbols as in Figure 7.
ment analysis is performed. We emphasize
the desirability of establishing a careful set of
El Nio indicators for every species used. This
clearly cannot always be tied into an opportu
nity for direct observation (as was the case
with the 1982-1983 event) but should always
involve careful and detailed integration Cif
growth increment andstable isotope tech
3. Ancillary information regarding sudden
and dramatic changes in taxonomic distribu
tion should be considered in any attempt to
recognize ancient El Nio events. Our obser
vations during the 1982-1983 El Nio sug
gest that such changes will most likely be
selective. Large clams (e.g . Mesodesma don
acium) and rock-dwelling gastropods (e.g.,
Fissurella spp.) might be preferentially af
fected. and an abrupt decline in abundance of
VOl. 2, NO. 3
00 ....-------...------,

- 0.8
Relative Growth Increment
Figure 12. &lBQ-growth increment distribution da
gram of Trachycardium procerum. Explanations and
symbols as in Figure 7.
these species in a particular stratum might be
significant. Sudden increases in thepercent
age of small mussels (especially Semimytilus
algosus and Brachiodontes purpuratus) and
the large gastropod Thais choco lata may sig
nal the presence ofa large scale El Nio evento
These species appeared to be minimally af
fected during the 1982-1983 evento The
small beach clam, Donax obesulus, recovered
quickly following the 1982-1983 El Nio and
its sudden abundance in a stratigraphic se
quence might also be indicative. Dramatic in
creases incertain motile species (e.g., the scal
lop Argopecten purpuratus) should al so be
This approach must be carefully applied, as
many taxic changes in shell middens may
indicate changes in resource exploitation un
related to El Nio events (e.g., shifts in har
vesting technologies or cultural preferences,
alterations in coastallandscapes, etc.). .
4. Perhaps the most serious potential con
straint upon application of these techniques
involves the nature ofthe midden record. Cer
tainly, the search for ancient El Nio events
will be most successful under conditions con
ducive to microstratigraphic sampling ofrela
tively undisturbed midden material. Shell
middens, however, are often culturally ds
rupted (although the midden material associ
ated with seasonally occupied base camps or
shellfish harvesting sites along coastal Peru
may be less culturally disturbed than shell
refuse heaps associated with more continu
ously occupied sites). Stratigraphic sampling
of midden material should be supplemented,
wherever possible, with sampling ofproximal
natural strata. Many ofthe sites along coastal
South America provide this opportunity (see
Rollins et al., 1986a), but even in these cases
care must be taken to avoid seriously time
averaged faunal assemblages. Ideal condi
tions for detection of ancient El Nio events
involved catastrophic burial; fortunately this
may commonly accompany an El Nio pertur
bation (e.g., beach ridge deposits associated
with El Nio flooding events as discussed by
Sandweiss, 1986). The temporal "window"
available to be recorded is admittedly small,
dependent upon the duration of the El Nio
event and the longevities of the bivalve spe
cies. Adequate conditions do exist, however,
in the fossil record, as documented by Clark
(1987) during growth line analysis of a
Pliocene scallop assemblage. The chances of
recovering a large scale El Nio event in the
ancient record are enhanced by the fact that
major El Nios have a high probability of
being closely spaced in time. Quinn et al.
(1978) assessed the probability of strong El
Nio events recurring within 7-8 years at 82
We gratefully acknowledge the research support of a
National Geographc Society grant and a S e ~ o r Faculty
grant from the Latn American Studies program at the
University of Pittsburgh. We profited from discussions
with Mr. Thomas DeVries of Oregon State University
and we are thankful for the drafting assistance of Mr.
Frank Benacquista.
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- ~ - - - ~ / '