Marine Geology - Elsevier Publishing Company, Amsterdam - Printed in The Netherlands

MARINE PALYNOLOGY: POSSIBILITIES, LIMITATIONS, PROBLEMS*

JOHAN J. GROOT1, 2 AND CATHARINA R. GROOT2

1 University of Delaware, Newark, Del. (U.S.A.)

2 Lamont Geological Observatory of Columbia University, Palisades, N.Y. (U.S.A.)

(Received March 25, 1966)

SUMMARY

In general, pollen and spores are present in the sediments of the deep ocean
basins, particularly in deposits of terrigenous origin. Pollen spectra of abyssal
sediments reflect the vegetation of adjacent land masses. Deposits of abyssal plains
and oceanic rises with relatively little local relief, situated in the temporate zones,
present the most favorable opportunities for obtaining a continuous history of
vegetational and climatic changes during the Quaternary.
The very slow rate of deposition in the ocean basins and the need to have
relatively large core samples in order to obtain a sufficient number of pollen grains
impose a limit on the detail with which reconstruction of vegetation and climate can
be accomplished. Occurrence of reworked plant microfossils presents difficulties
particularly where pre-Quaternary sediments are exposed on the continental shelf
or slope.

INTRODUCTION

Since the close of World War II knowledge of the geographic distribution and
thickness of various types of sediments in the ocean basins has increased tremendously
owing to extensive seismic-reflection work carried out by major oceanographic
institutions. In addition, much has been learned about the stratigraphy and rate of
deposition of sediments within reach of present coring techniques. This knowledge
is based on micro-paleontological work (e.g., ERICSONet al., 1961, 1964; RIEDELand
FUNNELL, 1964; HAYS, 1965), and on various methods of radiometric dating.
Palynology has also rapidly developed during the past 20 years. Until the last
war, it was nearly exclusively applied to the study of sediments of continental facies,
primarily organic deposits, but since then pollen and spores have been successfully
separated from marine sediments as well.

* Lamont Geological Observatory Contribution No.983.

Marine Geol., 4 (1966) 387-395

388 J . J . GROOT AN[) C. R. GROOI

The contemporaneous development of marine geology and palynology leads

logically to an attempt to apply pollen analysis to problems concerning the strati-
graphy of Cenozoic sediments in the ocean basins in addition to the "classical"
problems of vegetational and climatic changes of the Quaternary. Moreover, paly-
nology can make an unique contribution in the correlation of sediments of continental
and marine facies, including those of abyssal facies.
So far, very little has been published on the palynology of ocean basin sediments,
but we believe that a review of published and unpublished data available to us
warrants a provisional appraisal of the possibilities and limitations of "marine
palynology".

MARINE PALYNOLOGY

It is certainly no exaggeration to state that our knowledge of the history of

Quaternary climatic changes is based mainly on the results of pollen analysis of
peats and other continental deposits. Yet, there are two drawbacks connected with
this type of investigation:
(1) Most continental deposits contain unconformities, and consequently the
vegetational and climatic record obtained from them is a discontinuous one, and
often limited to a relatively short period of time. Although investigation of many
deposits, geographically and stratigraphically widely distributed, can minimize the
drawback of short-time or discontinuous records, some difficulties usually remain
in deciphering the history of the Quaternary.
(2) Pollen diagrams from continental deposits, and particularly from peats,
represent the vegetation of a rather small area adjacent to the site of deposition.
Thus, local environments have a significant influence on the composition of the pollen
diagrams, so that the latter are not entirely representative of the regional vegetation.
The same drawbacks and difficulties pertain, in some measure, to the study of
sediments of the continental shelf. Owing to Quaternary sea-level changes, deposition
on the shelf was probably discontinuous in many areas. Moreover, at least part of
the shelf is a rather high-energy environment where deposition of pollen grains is
unlikely. Sediments on the continental slope are probably largely unstable, and it does
not seem to be a promising environment for palynological work.
On the other hand, the deep-ocean basins appear to offer excellent opportuni-
ties, because:
(1) The chance of finding areas of continuous sedimentation, making it possible
to obtain continuous records of vegetational and climatic changes at least throughout
Quaternary time, appears to be good if cores are selected judiciously. Those areas
of the ocean floor which are not swept by rather strong bottom currents, and which
have so little relief as to make slumping unlikely, would appear to be the most favour-
able. Thus, we would expect that abyssal plains and those parts of oceanic rises
which have little relief are the most suitable areas from a topographic point of view.
Marine Geol., 4 (1966) 387-395
MARINE PALYNOLOGY~ POSSIBILITIES~ LIMITATIONS, PROBLEMS 389

(2) Generally, the ocean floor is a low-energy environment; such an environ-

ment is suitable to the deposition of pollen grains as well as fine-grained clastic sedi-
ments.
(3) Abyssal plains and oceanic rises are usually some distance from the conti-
nents, being separated from the latter by continental shelves, slopes and continental
rises. Thus the site of deposition is generally at considerable distance from the source
area of the pollen grains, and pollen diagrams from deep-sea sediments can be
expected to reflect the vegetation of large regions thus eliminating local environmental
conditions.
(4) Pollen grains can be considered as sedimentary particles. If we can accept,
for the moment, the hypothesis that pollen grains and fine-grained clastic particles
of terrigenous origin behave similarly, and are transported essentially in the same way,
then we may utilize knowledge of the provenance of pollen grains in interpreting the
provenance of the lutites in which they are found. This hypothesis is based on the
belief that water currents are a major agent of transportation of pollen and spores
to the deep ocean. This subject will be discussed later.
Factors that influence the occurrence of pollen grains in ocean basin sediments
are
(a) The type of sediment. Most favourable are lutites of terrigenous origin,
either because the lutites and the pollen grains were transported and deposited
together, or because preservation is better in lutites than in other types of sediments.
Less favourable are sediments with a high carbonate content, particularly Globigerina
ooze, and diatom ooze. Abundant biogenous material "dilutes" the terrigenous com-
ponent of the sediment, and therefore decreases the number of pollen grains per gram.
Where organic productivity is small as compared with the supply of terrigenous
material or where the ocean is deep enough to cause the solution of carbonates, pollen
grains are usually found. The suitability of turbidites for pollen analysis is uncertain at
present. The authors' initial expectation was that turbidite layers should be somewhat
older than the pelagic sediments on which they rest, because the sediment of which
the turbidite layers are composed was temporarily deposited on the continental slope
previous to its transportation to the abyssal plain. However, investigation of some
cores from the Argentine Basin failed to show any significant difference between
the pollen spectra of turbidites and those of other sediments (GRooT and GROOT,
1964). This seems to imply that there is no significant age difference between the diffe-
rent types of sediment in these cores and, consequently that the material incorporated
in the turbidite layers did not remain on the continental slope for a long time. Whether
this case is an exception rather than the rule can only be decided by study of additional
cores.
(b) Distance from shore. D. B. Williams (personal communication, 1966)
investigated the number of pollen grains per gram in North Atlantic bottom sediments
and showed a general decrease with distance from shore.
(c) Oceanic and atmospheric circulation undoubtedly play a role, but their
exact influence needs to be investigated. It has been demonstrated by MULLER (1959)
Marine GeoL, 4 (1966) 387-395
390 J . J . GROOT AND (~. R. GROOT

that pollen can be reasonably abundant in an area where atmospheric circulation

is adverse, and we found this to be true for sediments off the coast of central Chile
where westerly winds prevail. In such areas oceanic circulation must be mainly
responsible for the dispersion of pollen (GRooT and GROOT, 1966).
(d) The macro- and micro-relief of the ocean floor have probably as much
influence on the deposition of pollen grains as they do on the deposition of clays.
KORENEVA (1964) found high concentrations of pollen in depression sediments of
submarine relief.
As far as we can ascertain at the present, the most significant factor is the type
of sediment; it appears that the occurrence of muds of terrigenous origin is nearly
always indicative of the presence of pollen grains in excess of 20 per gram of sediment,
and often considerably more. The most favourable areas for marine palynological
work appear to be oceanic rises and abyssal plains where sediments of terrigenous
origin are predominant.
There are, of course, also some limitations and difficulties connected with this
type of investigation. The main limitation is that, with low rates of deposition prevail-
ing in the deep ocean, and rather large quantities of sediment needed to recover an
adequate number of pollen grains, each sample represents a considerable span of
time, say one thousand years or more; consequently it has not been possible so far
to trace minor vegetational changes and climatic fluctuations which often can be
demonstrated by pollen analysis of continental deposits. A second limitation, at least
from the botanist's point of view, is the impossibility to reconstruct ancient plant
communities, because deep-sea pollen spectra always reflect a regional vegetation,
or, as KORENEVA(1964) expressed it, the "spore and pollen spectra are more mixed,
thus giving an integrated picture of the vegetation".
A serious difficulty is the presence, in some areas, of pollen and spores reworked
from older deposits. These are apparently abundant in those areas where older for-
mations crop out on the sea floor, either on the continental shelf or on the continental
slope. For instance, pollen and spores in the Quaternary sediments of the Biscay
Abyssal Plain contain up to 50 ~/0 reworked material of Tertiary to Paleozoic age
(GRooT, 1963). Significant percentages of reworked pollen have also been found in
sediments of the continental rise off eastern North America. In both these examples
pre-Quaternary rocks are known to occur on the continental shelf or slope. Although
present-day rivers erode areas which could be the source of reworked pollen grains,
an investigation of the pollen and spores in the Delaware River and Bay indicates
that the amount of reworked material is considerably less than 1 ~, undoubtedly
because of the tremendous number of modern pollen grains that are transported in
the water (see also GROOT, 1966). Such a small percentage does not seriously interfere
with interpretation of Quaternary stratigraphy and climatic changes. It is, however,
not certain in how far the findings from this local example may be extrapolated to
other areas.
The problem of reworked pollen and spores is not very serious if they are of
Paleozoic or Mesozoic age. Usually, Mesozoic and older grains are morphologically
Marine Geol., 4 (1966) 387-395
MARINE PALYNOLOGY: POSSIBILITIES, LIMITATIONS, PROBLEMS 391

quite distinct from Cenozoic ones, and, in addition, they do not take Safranin O
stain in the same way as Cenozoic pollen do. The problem is more difficult with
Tertiary pollen, and the cautious investigator will undoubtedly make a study of
Tertiary grains that could possibly be present in Quaternary deposits. Autofluores-
cence studies such as developed by VAN GIJZEL (1961, 1967) may well be the solution
to this problem.

TRANSPORTATION OF POLLEN AND SPORES TO THE OCEAN FLOOR

It was suggested earlier that pollen and spores in ocean bottom sediments may
be indicative of the provenance of the lutites in which they are found, and this
hypothesis is based on the belief that water currents play a major role in the transpor-
tation of pollen grains and spores to the deep ocean, and that they are transported
together with fine-grained mineral matter of terrigenous origin.
In considering the possible validity of this hypothesis, some information con-
cerning the relative importance of wind and current transportation should be evalua-
ted. Unfortunately, the amount of data available on long-distance transportation
in the atmosphere is rather meager, and our knowledge of transportation by water
currents is even less.
Factual information can be summarized as follows:
(1) Data presented by ERDTMAN(1943) and DYAKOWSKA(1948) show that at
least 90 ~ of the pollen grains transported in the atmosphere are deposited in the
ocean within about 100 km from the shore. FAEGRI and IVERSEN(1964), discussing
atmospheric transportation, stated (p.37): "The distance 50-100 km thus forms a
natural limit of pollen dispersal. It is self-evident that the greatest quantities are
deposited long before this limit has been reached . . . . "Therefore, we can be reason-
ably certain that many, perhaps the great majority of the pollen grains found in
sediments 100 or more km offshore has been subjected to water transportation, and
it appears likely that the relative significance of water transportation increases with
distance from shore.
(2) The effectiveness of transportation of pollen grains by water currents was
clearly demonstrated by MULLER (1959) on the basis of the distribution of pollen
and spores in the sediments of the Orinoco delta, an area subject to the onshore
trade wind. CROSSand SHAEFER(1965) pointed out that stream transportation plays
an important role in plant microfossil distribution in the sediments of the Gulf of
California. Finally, ROSSIGNOL (1961), in her study of sediments off the coast of
Israel, found that many pollen and spores were transported by the Nile, and sub-
sequently by currents along the coast.
(3) Investigation of particles in suspension in Delaware Bay, in August 1964,
showed that the number of pollen and spores per gallon of water varied a great deal,
being high in the upper reaches of the estuary and during ebb tides, and being lowest
at the mouth of the Delaware Bay and during flood tides. The uniformity of the pollen
Marine Geol., 4 (1966) 387-395
392 J . J . GROOT AND C. R. GROO]

spectra and the presence, even at the bay mouth, of species growing more than 200
km from the sampling stations indicated that the pollen grain was derived from the
drainage basin of the Delaware River as a whole, and that local influence on the pollen
spectra was small. Furthermore, and perhaps of greatest interest, was that the number
of pollen grains in the Delaware samples was related to the amount of mineral
matter (GROOT, 1966); the number of pollen grains was about 20,000 per gram of
sediment. This was equally true for the upper and lower reaches of the estuary of
the Delaware River, and for flood and ebb tides. We should not draw the conclusion,
however, that there are no great variations in the pollen numbers per gram of sedi-
ment; during the winter months we would expect few pollen grains per gram of
sediment, and perhaps a greater number during the spring. Nevertheless, the data
suggest that in this case pollen grains and fine-grained clastic sediments were trans-
ported together and in the same manner.
(4) Data regarding the occurrence of suspended matter in the ocean shows that
most suspended matter is present in the lower few hundred metres of the water column
(EwING and THORNDIKE, 1965). Significant amounts have also been reported in near-
surface waters by LISITSYN(1961), but most of this is probably of biogenous origin.
On the basis of these data it is suggested that nine out of ten pollen grains
found in sediments in the deep ocean basins were transported by water currents,
near the surface or at depth, from nearshore areas (that is, from less than 100 km
from the coast) to their site of deposition. Therefore it appears safe to assume that fine-
grained clastic sediments transported in suspension across the continental shelf to
the deep ocean, will travel in company with the majority of the pollen grains on their
way to the same destination. Consequently the pollen grains should give some indi-
cation & t h e provenance of lutites.

MAGNITUDE OF POLLEN TRANSPORTATION TO THE OCEAN FLOOR

We may make some rough calculations as to the number of pollen that are
delivered to the ocean by streams on the east coast of North America. According
to GILLULY (1964), denudation in eastern North America resulting in suspended
sediment being transported to the ocean is in the order of 5 • 10-4 cm/year. Assuming
a specific gravity of the eroded rocks of 2.5, this produces about 12.5 tons/km 2
year. The total drainage area is about 1.8 • 106 km 2, and the amount of sediment
transported to the Atlantic Ocean from this area is in the order of 22.5 • 106 tons.
If we assume that the figure of 20,000 pollen grains/g is average for the pollen produ-
cing season, say during four or 5 months of the year, then the number of pollen
transported to the western North Atlantic by rivers is in the order of 15 • 1016/year.
Measurements of off-shore pollen rain from the atmosphere were made by
Hesselman (reported in ERDTMAN, 1943). He found that on two light vessels in the
Gulf of Bothnia 1620 and 880 pollen grains/cm 2 descended in the period 24 M a y -
26 June 1918; the vessels were 30 and 55 km off-shore. The order of magnitude of
Marine Geol., 4 (1966) 387-395
MARINEPALYNOLOGY:POSSIBILITIES,LIMITATIONS,PROBLEMS 393

the pollen rain in an area up to 100 km off-sh0re might be between 10z and 104
pollen/cm2 season. Off the east coast of North America, the seasonal pollen rain
would be approximately 15 • 1018; this is 100 times the amount computed for river-
transported pollen. Assuming (1) that the number of pollen grains delivered to the
western North Atlantic by atmospheric and river transportation is somewhere near
the figures mentioned above, that is in the order of magnitude of 1019/year, and (2)
that these are equally dispersed in sediments between the edge of the continental
shelf and the center of the Mid-Atlantic Ridge (an area of roughly 9 × 106 km 2
excluding the region of north of 50 degrees), then the pollen rain on the ocean bottom
should be approximately 100 grains/cm 2 year. Assuming an average sedimentation
rate of terrigenous lutite of 0.1 cm/10 z years, 1 cm z or about 2 g of terrigenous lutite
should contain 106 pollen grains or about 5 • 105 pollen grains/g. This compares with
an average of 20-40 grains actually found. Thus it appears that the "casualty rate"
is very high indeed. Nevertheless, pollen spectra from recent marine sediments do
reflect the vegetation of the adjacent land as was shown by KORENEVA(1964) in her
study of the sediments of the Sea of Ochotsk, and by GROOT and GROOT (1964)
and GROOT et al. (in press) in a study of the sediments of the Argentine Basin.
We must stress the highly speculative nature of this calculation and we believe
that it would be very worthwhile to make a long-term, quantitative investigation
of atmospheric and ocean current transportation of pollen grains.

CONCLUSIONS

(1) Pollen and spores in excess of 20/g of sediment are generally present in
samples of lutites of terrigenous origin even at considerable distances from the con-
tinents.
(2) Pollen analysis can be applied to sediments which do not lend themselves to
micropaleontological investigation, and it complements the work of ERICSON et al.
(1961, 1964).
(3) Pollen spectra from ocean bottom sediments reflect the main elements of
the vegetation of the adjacent land.
(4) Pollen spectra from oceanic rise and abyssal plain sediments in judiciously
chosen cores may provide an excellent opportunity to obtain continuous records of
vegetational changes occurring during the Quaternary.
(5) There are good indications that pollen grains and spores can be considered
as sedimentary particles indicative of the provenance of the lutites in which they are
found.
(6) Combination of the need for rather large samples and the prevailing low
rate of deposition in the deep ocean impose a limit on the detail with which vege-
tational history and climatic fluctuations can be reconstructed.
(7) The presence of reworked pollen in significant numbers appears to be a
problem particularly where polleniferous sediments crop out on the continental shelf
or slope. Auto fluorescence studies will probably alleviate this problem.
Marine Geol., 4 (1966) 387-395
394 J.J. GROO[ AND (. R, GROOI

ACKNOWLEDGEMENTS

We are indebted to Professor Maurice Ewing, Director of the Lamont Geolo-

gical Observatory, for making available cores collected under his direction in large
part aboard the "Vema", with the financial support of the Office of Naval Research
(contract No. 266 (48)). We thank him and David Ericson for critically reading the
manuscript, and Charles Fray and Roy Capo of the Lamont Core Laboratory for
assistance during the sampling of the cores. Sampling in the estuary of the Delaware
River was financially supported by the University of Delaware Research Foundation.
This paper was written during the tenure of a John Simon Guggenheim Fellowship
by the senior author.

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Marine GeoL, 4 (1966) 387-395