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SUMMARY
This article deals with the most important aspects of nearly twenty years of
intensive study of the pollen-and-spore content of Tertiary sediments in some parts
of tropical South America, Africa and Asia.
For a proper evaluation, the character of the data, including the selection
and preparation of the samples, the diversity of previous recording and the statis-
tically achieved uniformity in presentation of these basic data needs a full expo-
sition, given in the introduction. This is directly followed by an explanation of the
process of elimination of all stratigraphically unimportant species. The resulting
interpretation of climatic and topographical influences on the dispersal of pollen
and spores is illustrated with examples from the fossil record. The disturbing
effect of redeposition forms a problem, which in some cases can be solved. Now
that the main ways of dispersal of pollen and spores are understood, the charac-
teristics of the three major depositional environments can be distinguished by
purely statistical analysis, without necessarily having any botanical information
from probably related Recent plant species. Additionally botany and palae-
ontology may bring supporting evidence. This many-sided approach leads to the
discrimination between local and regional features of environmental or time-
stratigraphical significance which is needed for the evaluation of long-distance
correlation.
As a result the marker species can be classified into: (1) a restricted number
of pantropical marker species; (2) a larger number of marker species which oc-
curred in both the South American and west African regions, tropical today
(transatlantic distribution); and (3) a still greater quantity of species which are of
significance only within a single botanical province (intracontinental distribution).
Thus a broad stratigraphical framework on a pantropical scale is established,
which may be further subdivided regionally. These three systems of subzonation
are compared with independent zoopalaeontological time-stratigraphical cor-
relation and discussed in great detail, with special emphasis on the Carribean data.
INTRODUCTION
This paper presents some of the results obtained in twenty years of palynolo-
gical investigation of Tertiary sediments from tropical areas by companies of the
Royal Dutch/Shell Group. The primary purpose of this investigation was to
arrive at a better interpretation of the stratigraphy of those Tertiary sedimentary
basins where other means of correlation failed. Of course it is not possible in
this paper to do more than present an outline of the large amount of work done
and to select for discussion a few topics which appear to be of more general
interest. Documentation will also be restricted to selected examples.
The restriction of the subject of this paper to the Tertiary of tropical areas
is due to a combination of factors. Firstly, the tropics represent a natural geobotan-
ical unit. Secondly, palynological research by Shell was initially mainly concerned
with the Tertiary sediments of northern South America, Nigeria and Borneo.
Thirdly, the amount of detailed information on the Upper Cretaceous palynolo-
1 In this connection we should like to express our gratitude for the facilities extended to us by
Dr. T. van der H a m m e n (Amsterdam) and his staff, which enabled us to study the type collections
not only of already described species but also of types to be described in future publications.
where a : the amount, that is the number of grains of the single species observed
in the old sample; N : the sum of the amounts of all species, that is the total
number of grains or pollen sum observed in the old sample; M : the sum of the
amounts of all species in the new sample.
Depending on the accuracy required by the palynological investigation and
the time and sample material available, the value of M may be taken larger or
smaller than 100. For technical reasons related to the type of computer at present
in use for palynological data processing, the value of M is here taken to be 88.
This conforms approximately to the current sample size of 70-I00 specimens of
the selected species. The advantage of such a relatively small sample size lies mainly
in the possibility of examining large numbers of samples within a short time, which
is essential for routine work, both stratigraphically and statistically, and is far
more informative than a few samples with large pollen sums.
In general the counting of the marker species takes place in two phases.
First, an analysis is made of the whole flora in which the dominant species like
Rhizophora are also counted. The second part of the microscopical investigation
is concerned only with the more rare but important markers, among which, e.g.
Asteraceae (Compositae). To save time dominant species are no longer counted,
but statistically it is not permissible to multiply the amount counted to get the
corresponding figure. Therefore, the probability of re-observation of, for instance,
Rhizophora pollen is calculated from the first pollen sum, but the probability of
re-observation of the rare markers like Echitricolporites spinosus from the second
pollen sum.
Example." I - - c o u n t o f t h e w h o l e f l o r a , i n c l u d i n g , i n t e r alia, Rhizophora -- 100 g r a i n s .
II : c o u n t o f t h e s p e c i a l s e l e c t i o n , e x c l u d i n g Rhizophora : 84 g r a i n s .
A m o u n t o f Rhizophora in ' T ' : 8 0 g r a i n s , p r o b a b i l i t y o f r e - o b s e r v a t i o n = ! - - (1 - -
8 0 / 1 0 0 ) ss : 1.00.
A m o u n t o f Echitricolporites sphwsus in " I " = l g r a i n , p r o b a b i l i t y o f r e - o b s e r v a t i o n =
1 - - (I - - 1/100) ss == 0 . 5 9 0 .
A m o u n t o f Echitricolporites spinosus in " l I " = 4 g r a i n s , p r o b a b i l i t y o f r e - o b s e r v a t i o n =
1 - - ( I - - 4 / 8 4 ) ss = 0 . 9 8 6 .
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of re-observation of this species in I is of little interest and is not recorded on the distribution
charts presented. This example also shows the importance of further counting of the selected
species.
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G E O L O G I C A L DATA
100J20'
120 140 ~ E El El
140.160' I~ El I~
160-180~ /
220 240' ~
240-260~ ~
260 280'
280300' .3
300320' 0!
340.360' ~
360 380' E ! El
I
380.400' C IO
400-420' I~ ~ Q
420.440' El
445-448' * / / ,/" *
504 $24' ~ El El
524.544' *
640 660 r * El El
396 140; 0 i0
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460147 *
525.153, /'1//
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570-15~(
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830-185( ~ ~)
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064-208a
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2177'
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~ z ~ ~ N ZONES ZONES ZONES
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MAGDALENEHSIS
PROXAPERTITES
OPERCULATUS
PROTEAC[DITES
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BUTTINIA ANDREEVI
PROIEACIDITES DEHAANI
STEPHANOCOLPITE$COSTA/US
FOVEOERILETESNARGARITAE
(~ " LONGAPERIiTES VANEENDENBURGI
~. RETIDJPORITESMAGDALENENSIS
~) '%. %`. ~) 0 '%. %` ~. 0 ECHJTRIPORITESTRIAHGULIFORMIS
~. (~ (~ %.. ~ ~. ~. PROXAPRRnTESOPERCULATUS
SPINIZONOCOLPITESGROUP
'%. " '%. 0 \ " 0 0 %`. CTENOLOPHONIDITES COSTATUS
%~. ~ ~ (~ RETISTEPHANOCOLPITESWFLLIAMSI
GENMASTEPHANO'COLPITESGENMATUS
[ - J ANACOLOSIDITRSCE LUTEOIDE$
j ".% ~ %, ~. ~,~1%` . O %` PROXAPERTITESCURSUS ~n
BOMBACACIDITES AHNA~ 0
t CTENOLOPHONIDITE$LISAMAE
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J %` %` Q " %` 0 (~) \ %. ~ ~ '%. ~) Q ~ RETIBREVIIRICOLPITESIRIANGULATUS
j %` * ~ 0 " (~ (~ 0 ~11 ~) ~) %, (~ 0 "%` SIRIATRICOLPITESCATATUNBUS
RETIIRICOLPORITESIRREGULARIS r-
J 0 '%. %` PSILATRICOLPORITESCRASSUS ~-~
7
I ~. %` O \ %` %. (~ %, ' ~. O \ MONOPORITESANNULATUS
PSILATRICOEPORITESOPRRCULATUS ~. r--
J MARGOCOLPORII~S VANWIIH1 (~
J RETITRICOLPORITESGUIANENSIS
j CICATRICOSISPORnESDOROGENSIS .-~ Z
PERISYNCOLPORITESPOKORNYI 0 0
J %` E) (~ ~ ~E) (~ (~ VERRUCATOSPORITES
USMENSIS rn
J %` PERFOTRICOEPITESDIGITAIUS "~
. . . . . . . . . . . . . . PACHYDERMIIESDJEDERIXI )~"
I ZONOCOSTITES RAMONAE ~ "~I
1 ECHIP~RIPORIIESESTELAE ~:) ~0
J ALNIPOLLENIIES VERUS
J FLORSCHUETZIAIRILOBATA ..~ ~1~
JANDUFOURIA SEAMROGIFORMIS t~ --I
[ - - VERRUERICOLPORITES
ROTUNDIPORIS
J MAGNASTRIATIIES HOWARDI ~ --
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J FLORSCHUEIZIA LEVtPOLI ~>
J CRASSORETITRILETESVANRAADSHOOVENI ,~
J MULTIMARGINITES VANDERHAMMENI -0
GRIMSDALEAMAGNACLAVA[A "1-
J FLORSEHUETZIAMERIDIONALIS
JI ECHITRICOLPORIIESSPINOSUS
I STRIASYNCOLPIIESZWAARDI
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J ECHIIRICOLPORIIESMCNEILLYi r~
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I ~ ~ o
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pollen suspension entering the sea. Also, the largest number of species will be
present in the fluviomarine sediments.
In the second case the composition of the pollen load is quite differently
affected. First of all sea currents act as slow transporting agents over wide fronts
with relatively less turbulence further offshore; the result is gradual settling and
deposition of pollen grains and spores, accompanied by size sorting. Wind-trans-
ported pollen may be added and this may considerably modify the composition
of the pollen load in areas far offshore. Another factor about which little is known
is selective corrosion of pollen and spores during seaborne transport.
One final complication has to be evaluated and this is the probability that
not all pollen is travelling in a single move from anther to final resting place, but
that temporary entrapment in sediments is followed by subsequent erosion and
that renewed transport takes place. This recycling of pollen grains and spores may
take place within what is geologically speaking an insignificant period of time, as
is the case in the average delta, where meandering streams may erode sediments
deposited only a short while before. Such recycling, of course, only increases the
degree of mixing of the pollen load eventually supplied to the sea and does not
affect stratigraphical distribution patterns to any appreciable degree. However,
when the time lag between deposition and recycling becomes greater, serious
difficulties may arise, culminating when older pollen-bearing formations are eroded
in the source area of a river system.
When a clear difference in preservation between such older reworked pollen
grains and spores and autochthonous ones is present, they can be easily excluded
from the pollen sum, but unfortunately this is usually the exception rather than
the rule. Generally there is no clear-cut difference, and the presence of many
slightly different corrosion patterns in one single sample may be the only clue
indicating the presence of several generations of reworked pollen grains and spores.
In such cases a detailed knowledge of the basic floral succession in the area of
investigation may enable one to spot the scattered presence of anomalous pollen
associations and to recognize them as reworked assemblages. Here general geolog-
ical considerations, indicating the probable origin of the sediment particles have
also to be taken into account. Recently VAN GIJZEL (1961) has demonstrated that
fluorescence microscopy may be able to discriminate between autochthonous and
reworked pollen.
In the Palaeogene sediments of western Venezuela, for instance, reworked
pollen was scarce because the bulk of the sediments deposited during that time had
been derived from the Guiana shield, where no pollen-bearing sediments were
eroded. In younger sediments the occurrence of reworked pollen of a Palaeogene
age proved to be clearly related to distinct unconformities.
In contrast, in the Neogene sediments of northwestern Borneo no such
clear-cut unconformities exist and since the sediments had been recycled many
times during the course of the Tertiary, a rather diffuse distribution of reworked
Examples
The practical application of the above principles will now be illustrated with
a few examples.
The first case refers to base Monoporites annulatus Zone in northern South
America. Within the Eocene of western Venezuela, which was the first area inves-
tigated, two floral boundaries were apparent in approximately the same strati-
graphical interval: aal well-defined increase in Echitriporites trianguliJormis and,
slightly higher, a less conspicuous base of Monoporites annulatus. Later the absence
ofEchitriporites trianguliformis in the Eocene of Colombia was noted. This absence
could not be attributed to a stratigraphical hiatus or to a different climate, since
the remainder of the floral assemblage proved to be essentially similar. However,
the Colombian Eocene sediments are more terrestrial than those in Venezuela, and
it became clear that Echitriporites trianguliformis was derived from a coastal plant
with mainly waterborne dispersal of its pollen in seaward direction. This reduced
its value for regional correlation and, since Monoporites annulatus appeared to
have a wider distribution, because it was derived from a more inland environment,
its increase in the Early Eocene both in marine and more terrestrial deposits is
preferred as the regionally valid time-stratigraphical correlative horizon. The
marked increase in Echitriporites trianguliformis retains, however, its practical
correlative value within the restricted area of the Lower Eocene in the Maracaibo
basin.
In the case of the top in the occurrence of Proteacidites dehaani (Upper
Cretaceous) it was soon apparent that this event could be recognized both in
marine deposits in the northern Maracaibo basin and in alluvial plain deposits in
Colombia and the southern Maracaibo basin. Regional time-stratigraphical value
could, therefore, be assumed for this floral boundary.
In both these examples botanical identification of the pollen species was
impossible or did not provide sufficient evidence to permit an independent check
on the environmental requirements of the parent plants. The method outlined
above was the only feasible one for the recognition of time-stratigraphical value.
In the examples from the Neogene, described below, on the other hand,
this sort of conclusion is indeed supported by direct botanical evidence.
Z ~ _~
z ~ ~ ~ ~ ~ ~ ~ ~ ~
~6~o' / . .
z8~6, / . /
~;'~' 0
2931' .~ .
==- ~oo~, I /
3140' " ! "3
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3475' .
3723' / O O /
m ~ sgso, . . ..
3980' / O O /
m%, O / ~ . .
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o oo, . . . / . . /
45,,0, / /0 - . /
4650, / " 0
4725' " Oi / /
4805'
CASSIGERINELLOITA 4880'
AMEKII ZONE
SOBS, / / / /
SUBDIVISION
~_~ _
Oz
>~ ~ ~1 ~ z ~ ! CARIBBEAN ATLANTIC PANTROPICAL
o ~ ZONES ZONES ZONES
az
VERRUTRICOLPORITES
RO1UNDIPOR~S
NAGHASTRiATITES
HOWARDI
CKCATRICOSISPORITES
DOROGENSIS
VERRUCATOSPORITES
USMENSIS
MONOPORITES
ANNULATUS
z
@ O - k
\ \ @ Q O O O . ~ @ O O O O O O O O ' O - @ @ O O O O @ O @
0 0 0 @ 0 0 0 @ 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Q O Q O \ .
0 0 0 0 0 0 0 0 0 0 0 9 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 \ - . 0 \ 0 0 0 - 0 0 0
\
O O O @ \ @ @ O O O O O O O O O O O 0 0 0 0 0 Q O O O 0 \ 0 0 0
\
GROUP
BENIN FORMATION
~ _
c~
C
Z r-
0 0
BUITINIA ANDREEVI
PROIEACIDITESDEHAANI
STEPHANOCOLPITESCOS~'ATUS
FOVEOTRtLETESHAR~ARITAE
LONGAPERTITESVANEENDENBURGI
RETIDIPORITESMAGDALENENSIS
ECHITRIPORII"ESTRIANGULFFORMIS
PROXAPERTITESOPERCULATUS
SPINIZONOCOLPITESGROUP
0 0 ~ CTENOLOPHONIDITES COSTATUS
RETISTEPHANOCOLPITESWrLLIAMSI
GEHNASTEPHANOCOLPITES GEMMATUS
ANACOLOSIDITESCF [UTEOIDES
rn
PROXAPERTITESCURSUS r-
BONBACACIDITESANNAE
CTENOLOPHONIDITESLISAMAE
FOVEOTRICOLPITESPERFORATUS
RETIBREVITRICOLPITESTRIANGULATUS
\ ~ STRIATRICOLPITES CATATUMBUS (~
REI'ITRICOLPORITESIRREGULARIS ~r-
(~ ~ '~. PSILATRICOLPORITES CRASSUS m
z )"
" 0 %. ' 0 ~. NONOPORITES ANNULATUS
'~, O ~ - PSILATRICOLPORITES OPERCULATUS 7'~ r--
MAEGOCOLPORiTESVANWUHEI (~ "-~
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CICATRICOSISPORITESDOROGENSIS z
PERISYNEOLPORITESPOKORNYI C) 0
VERRUCATOSPORfTESUSNENSJS m
PERFOTRICOLP~TESDIGITATUS "~
%, G PACHYDERMITES DIEDERIXI ~. IU~
ZONOCOSTITES RAMONAE ~ ""4
ECNIPERIPORITESESTELAE m ~XJ
ALNIPOLLENITESVERUS
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0 VERRUTRICOLPORITES ROTUNDIPORIS LJ~ --
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FLORSCHUETZIALEVIPOLI
"%. ~ \ CRASSORETITRILETES VANRAADSHOOVENt ~1>
MULTIMARGINlIES VANDERHAMNENI
GRIMSDALEAMAGNACLAVATA "1"
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ECHITRICOLPORITESSPINOSUS
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GEOLOGICAL DATA PALYNO-STRATIGR
ROCK.STRATIGRAPHICAL FAUNAL DATA SELECTED P O L L E N A N D SPORE M A R K E R SPECIES
UNITS
Z ~ i
- ~.,~ ~,
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3352' / ~ /
4~o, / / /
4376'
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6634' 'D ,
66S2' 0 .~
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712e,
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SUBDIVISION
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VANRAAOSHOOVENI
I GEOLOGICAL PALYNOLC
SUBDIVISION
zN
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z_,--
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UPPER SPINOSUS
18
CRA$SORETITRILETE$
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VERRUTRICOLP(
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26
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49
LOWER RETIBRE~TRI(O
TRIANGULAT
54
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OPERCULATUS MAGDALEHEI
DANIAN
65
~
-- PROTEACID|
Fig.15 (pp.230-232). Range chart, z~ MAASTRICHTIAN DEHAAN
Ln
70
~GICAL SUBDIVISION I SELECTE
C I CARIBBEAN I BORNEO
ZONES ZONES
ALNIPOLLENITES
VERUS
ECHITRICOLPORITES
MCNEILLYI
FLORSCHUETZIA
NERWIONALIS
PACHYDERM~ES
DIEDERIXI
GRIMSDALEA
--
._MA~~_ACL._AV_.AT__A
-- LEYIPOLI
PSILADIPORffES
MINIMUS
,R~TES
)ANDUFOURIA
SEANROGtfORM~S
I FLORSCHUETZIA
~RITES
S I TRILOBATA
GUIANENSIS
OPERCULATUS
- ~I~r'~c'G'L~t~E~ "~
.PITES
US
FOVEOTREOLPITES
PERFORATUS
ES I CTEHOLOPHONIDITES
ISIS LISAMAE
FOVEOTRILETES
MARGARtTAE
D POLLEN AND SPORE MARKER SPECIES
In the Oligo-Miocene the first increase in the Rhizophora pollen type (Zono-
costites ramonae) can only be reliably observed in coastal and offshore marine
sediments, while the approximately contemporaneous first increase in the Ceratop-
teris type (Magnastriatites howardi) can be detected in terrestrial environments as
well. However, the small fresh-water ferns of the genus Ceratopteris can only
flourish in open vegetation and will hardly occur under a closed forest canopy,
which restricts the dispersal of the spores to water transport mainly. Ceratopteris-
type spores thus occur more regularly in inland environments, where Rhizophora
pollen can be extremely rare, and therefore their oldest occurrence has been taken
as the more reliable time-stratigraphical horizon. In practice of course both floral
changes support each other and are used in conjunction.
Another example from the Neogene of western Venezuela concerns the
recognition within the same body of strata of three sets of correlation lines:
(1) A topographically-influenced succession of mangrove dominance alter-
nating with more inland vegetation.
(2) A subregional succession, probably of a climatological nature.
(3) A top occurrence of Cicatricosisporites dorogensis.
The correlation lines derived from (1) cross at an angle those from (2),
while none of these were observed in the adjoining fully marine succession. How-
ever, the top occurrence of Cicatricosisporites dorogensis proved to be recognizable
regardless of environmental influence, and thus was judged to be of major time-
stratigraphical significance.
In general the principle may be formulated that any palynological change
which is paralleled by a facies change is of questionable time-stratigraphical value.
if, on the other hand, the pattern of palynological changes crosses the environ-
mental correlation lines derived from lithology or benthonic faunal distribution,
then that pattern is likely to be of time-stratigraphical value at least within the
area of study.
Summarizing twenty years of experience, it may be stated that for most cases
tools are available which allow discrimination between local and regional events
of time-stratigraphical value on the one hand and effects caused by time-crossing
events such as transgressions and regressions on the other. This information then
allows the establishment of a zonation adapted to the particular stratigraphical
problem at hand. For short distances and short time intervals the effects of striking
topographical events may provide the correlative framework; for correlation over
larger distances climatological changes will produce the best means, while on an
intercontinental scale correlation on evolutionary change appears to be the only
reliable method.
In general, however, the larger the distance, the more difficult it becomes
to define and trace sharp floral boundaries. In particular boundaries defined on
evolutionary change are notoriously hard to place accurately in thick continuously-
deposited sediments. This often poses a practical problem, since the geologist
generally requires correlation lines, while the palynological events on which the
zonation is based are by nature mostly gradual. In fact any sharp floristic boundary
is more probably indicative of a hidden unconformity than of anything else.
The zones recognized in this paper are biostratigraphical units in the sense
of the AMERICAN COMMISSION ON STRATIGRAPHIC NOMENCLATURE (1961). They
are, therefore, not a priori to be considered as time-stratigraphical units and in
fact some of the boundaries delimiting the zones probably are not contempora-
neous over large distances. However, within the areas in which the zones are
recognized, their succession is identical in all sections investigated.
According to their lateral extent, the zones are grouped as follows: (1)
pantropical zones; (2) transatlantic zones; (3) intracontinental zones.
The definition and description of the zones will be documented by a restricted
number of type sections, mainly taken from northern South America (Fig.l-14).
The range chart (Fig.15) illustrates the ranges of all species discussed, based on
evidence from a much larger number of sections. The independent dating of the
zones is discussed in the section "Independent dating".
Pantropical zones
Intracontinental zones
Caribbean area
In the Caribbean area the transatlantic Retidiporites magdalenensisZone can
be subdivided into three units. The oldest unit is the Foveotriletes margaritae
Zone, which is characterized by the co-occurrence of frequent Stephanocolpites
costatus, Foveotriletes margaritae, Longapertites vaneendenburgi, and Gemma-
stephanocolpites gemmatus, and by the absence of Bombacacidites annae and
Ctenolophonidites lisamae.
The boundary with the overlying Ctenolophonidites lisamae Zone is taken
at the first occurrence of Ctenolophonidites lisamae and Bombacacidites annae, and
this zone is further characterized by the regular presence of Gemmastephanoeolpites
gemmatus, diminishing quantities of Foveotriletes margaritae, and the regular
presence of Proxapertites cursus.
The base of the youngest Foveotricolpites perforatus Zone is defined by the
first occurrence of Foveotricolpites perforatus, which species is restricted to the
zone. Bombacaeidites annae and Ctenolophonidites lisamae are frequent in this
zone, while Stephanocolpites costatus, Foveotriletes margaritae and Gemmaste-
phanocolpites gemmatus have virtually disappeared.
This subdivision is clearly visible in the Rubio-road section (Venezuela)
(Fig.3).
The Monoporites annulatus Zone can be subdivided in the Caribbean area
into four units. The boundary between the Psilatricolporites crassus Zone and
the overlying Psilatricolpites operculatus Zone is taken at the lowest occurrence
of Psilatricolpites operculatus, as visible in Icotea-1 (Venezuela), (Fig.4) and the
Prevenci6n section (Venezuela) (Fig.5).
The next higher boundary between the Psilatrieolpites opereulatus Zone and
the Retitricolporites guianensis Zone is placed at the base regular occurrence of
Retitricolporites guianensis. This boundary is present in the Prevenci6n section
(Venezuela) (Fig.5).
The Verrutricolporites rotundiporis Zone can be subdivided into the Jan-
dufouria seamrogiformis Zone and the overlying Psiladiporites minimus Zone. The
boundary between these two zones is marked by the base of the regular occurrence
of Psiladiporites minimus, as is visible in B-188 (Venezuela) (Fig.6). The Cras-
soretitriletes vanraadshooveni Zone can be subdivided in northern South America
into the Multimarginites vanderhammeni Zone and the overlying Grimsdalea magna-
clavata Zone. The boundary between these two zones is marked by the base of the
occurrence of Grimsdalea magnaclavata. This is clearly visible in B-188 (Venezuela)
(Fig.6).
Borneo
In Borneo a local subdivision can be established, based exclusively on the
evolutionary development within the genus Florschuetzia (Fig.16).
The lowermost zone, FIorschuetzia trilobata Zone, is characterized by the
presence of F. trilobata and the absence of younger forms. The zone probably
covers the upper part of the Verrucatosporites usmensis Zone and the lower part
of the Magnastriatites howardi Zone, but the stratigraphical position of its base
has still to be determined precisely.
The first development of Florsehuetzia semilobata and F. levipoli marks the
base of the next higher Florschuetzia levipoli Zone, which coincides approximately
with the weakly defined base of the Crassoretitriletes vanraadshooveni Zone. In
this zone FIorsehuetzia trilobata decreases in numbers, while F. semilobata dis-
appears. The average size of F. levipoli increases also from 30 /z to 35 /~. The
overlying Florschuetzia meridionalis Zone is characterized by the regular presence
of F. levipoli and, in increasing quantities, of F. meridionalis, while the average
size of both species shows further increases. Florschuetzia trilobata is still present
in the lower part of this zone, but disappears soon.
The boundary between the Florschuetzia levipoli and Florschuetzia meridio-
nalis Zones coincides approximately with the base of the Echitricolporites spinosus
Zone, and the first regular occurrence of E. spinosus and Florschuetzia meridionalis
can be taken as a criterion for its recognition.
Both species have the disadvantage that in the very thick Miocene sediments
of northwestern Borneo, their increase in number is gradual, which makes it
impossible to recognize a sharp boundary.
INDEPENDENT DATING
In western Venezuela and Colombia this zone is associated with the smaller
Foraminifera GIobotruncana gansseri, G. lapparenti, G. stuarti, Guembelitria ere-
tacea, Siphogeneroides bramletti, and the ammonite Sphenodiscus sp. In Nigeria
it is associated with the smaller Foraminifera Bolivina afra, Rugoglobigerina rugosa,
and striate Heterohelix spp., with the ammonites Didymoceras sp., Libyoceras
ismaeli and Sphenodiscus sp., and with the lamellibranchiat Inoeeramus sp.
This faunal list clearly indicates a Maastrichtian age. So far no evidence of
an older age has turned up, but since the lower limit of the Proxapertites opereu-
latus Zone has not been properly studied, the lower part of the Proteacidites
dehaani Zone may of course be older than Maastrichtian.
Unfortunately, both in Nigeria and in the Caribbean area, the top of the
Proteacidites dehaani Zone is present in a predominantly coastal facies without
In Colombia the lower part of this zone is associated with the larger Fora-
minifer Actinosiphon barbadensis, indicating a Paleocene age. In Nigeria the lower
part contains the smaller Foraminifera Globorotalia velascoensis, and G. acuta,
also indicative of Paleocene, while in the higher part the smaller Foraminifera
GIoborotaliaformosa and Globorotalia rex, which are markers for Stolk's Globoro-
talia.formosa range zone, indicate an Early Eocene age.
In view of the fact that in Venezuela the overlying Psilatricolporites crassus
Zone carries already a Middle Eocene fauna, while in Nigeria the overlying Mono-
porites annulatus Zone is associated with a Lower-Middle Eocene fauna, it is
suggested that the Retibrevitricolpites triangulatus Zone ranges from Late Paleocene
to Early Eocene in age.
In Venezuela the lower part of this zone is associated with the larger Fora-
minifera Linderinafloridensis, Helicostegina gyralis and Lepidocyclina sp. A, while
the upper part was found to contain Helicolepidina spiralis form C. According to
VAN RAAOSHOOVEN (1951), these faunal associations indicate an early Middle
and a late Middle Eocene age respectively.
Since the lowermost part of the overlying zone in Venezuela still is of late
Middle Eocene age, this boundary must be situated in the Middle Eocene.
This zone is associated in the Caribbean area with the smaller Foraminifera
Catapsydrax stainforthii and Globigerinatella insueta, and approximately covers
the Catapsydrax dissimilis, C. stainforthi and part of the Globigerinatella insueta
Zones of Bolli (Lower Miocene).
This zone was found to contain the smaller Foraminifera Globorotalia fohsi
robusta, G. mayeri and G. menardii and thus covers approximately the Globorotalia
fohsi robusta, Globorotalia mayeri and Globorotalia menardii Zones of Bolli. The
boundary between Lower and Middle Miocene would thus be present within the
zone.
The age range of this Bornean zone is determined by the presence, in the
lower part, of the smaller Foraminifer Globigerinatella insueta, indicative of Bolli's
Catapsydrax stainforthi and Globigerinatella insueta Zones, in the middle part, of
the larger Foraminifer Flosculinella bontangensis, indicative of Tertiary f 1, and
of the smaller Foraminifera Globorotalia fohsi barisanensis, G. foshi fohsi and
Orbulina universa, indicative of Bolli's Globorotalia fohsi foshi Zone, and, in the
upper part, of Globorotalia fohsi lobata, G. fohsi robusta and G. menardii, indicative
of Bolli's Globorotalia fohsi robusta and Globorotalia menardii Zones.
In Borneo the lower part of this zone carries the smaller Foraminifer Globo-
rotalia menardii, indicating an age younger than Bolli's Globorotalia fohsi lobata
STRATIGRAPHICAL APPLICATION
F 1~0 KM I 115 KM I 9(
C. DOROGENS,SZONE ~ I
V. U S M E N S I 5 Z O N E ) v. USt
........ = (A) P. C R A S S U S Z O N E
R. T R I A N G U L A T U 5 ZONE R . = ~.[
}~i;[i~:::..\ \ /
\\"
.......... ::i~-~:.iiii.-:.:iiiiii?::i;.-"iisi.:.-"~;i~iis
VERIICAL SCALE
:iiiiiiiiiiiiiiiiii~!iiiiiiiiiiiiiiii
:: ?iiiiiiiiiiiiiiiiiiiiiiiiiiiiii
METRES FEET :~i~i!i i i:ili~i i l!i !~ii i!i i i i;ili!i~i i i
0 - 0
~ii i!i:i!i!ililili!i!i!i~i~:i!i!i~i:i i i!i!i~i~i~i~i
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500 - ":::::::::::::: .-.
i!:::i:i:i:!:!:!::~'o:!!i!iiiiiii
2oo0 ::F:!:i?iiiii:?!F!i:~,~:??!ii!i
m 3000 :::::::::::::::::::::
*00o - : ::2:::::
' 4000
1500 -
5OO0
4 5 6 7 8 9
o ~_
OGENSIS ZONE
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::~:>.::::::::::::::::::::::::::::::::::::::::t o / !t::~:~t:~.:~:..-,~:!~::~.~.~!~.-:.*,::!::~:~.,::#:i
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~ ' NENsis"ZO'NEi)iiii?
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................................. . . . . . . . . . . . . . . . . ===================================================================
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. . . . . :.:.:-:-:-::.::-:-
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i:i~::.. ~t..*:~~~~!~ t,*'.'~,~!=============================
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:-~iii~~i~.~'~-~'.;.!~. ~ i :::::::::I:~........................~ ~ : . : : : : ~ I .......... .L
:, ::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::: ::::::::::::::::::::: (3)
:~F::.-::',:.~.~.~.F:$~.:'...~:~).~: :::::::::::::::
VENEZUELA
' '%!ii
I 60 KM I ISO KM I 140 KM I
7 P. CRASSI
R. TRIANGL
i / ~ .............i!iiiii~i~"
/
/ / :.....!7'
/ / , :~:~....
-- :: ~ MARINE
:.>:
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ii!i77
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' ~ ..Y
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iS ZONE
JSI5 ZONE
JS ZONE
fLATUS ZONE
ENENSIS ZONE
N I
Fig.17 (pp.249-252). Stratigraphical section I.
1
CHAFURRAY-3
<
G. MAGNACLAVATA ZONE
M. VANDERHAMMENI ZONE
ROTUNDIPORUS ZONE
C. DOROGENSIS ZONE
V. USMENSIS ZONE
.....
VERTICAL SCALE
METRES FEET
0 - - 0
* 1000
500
2000
3000
1000
4000
1500
5OOO
VORAGINE-1 RIO
0 KM { 375 KM
::::::::::::::::::::::::::::::::::::: ....................................................`.................................
: :i:i:i:i:i:i:i:i:i:i:i:i:i:i:i:].:. :::::: :::::::::: ::;:::::;:::: ::::: :::::::::::::::::::::: :::: ::::::::::::::::::::::::::::::: ::: ;:::::::; :;:;:::;::::::::;:: :::::::::::::::::::: ::::::::::::::::::::::::::::::: :;::::;:::: :;:::::: :::::::::::: :::: :::::::::2;:: :;2;: ::::::::::::::::::::::::::::::: ;:::::;:::::::::2 :::::::::2 :::::::::::::::::::::::::::::::
::::::::::::::::::::::::::::::::::::: ~:::~:~::::~:::::::::::::::::::::::::::::::~:~::~:~:]:~:~:~::::::::~:~:~:~:::~:~:::::::::::::::::::::::~:~:~:~::~:::~::::::::::::::::::::::::::::::::::::~:~:~:~:~:~:~:::~:~::::~:::::::::::::::::::::::::::::::::~;~;;;~;;
================================= :::::::::::::::::: ::;:::::::::::;:;; ;;:::::::::::::::::::::::::::::::: :::::::::::: :::: ;:;:::;:::::::;;:::: ;:;;:::;::: ;:;:::::;:::::: ::::::::: :::::;:::::::::;::: ::::;::; ;;;:;:;;:::;:: :;:::;:::;:: :::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::: ::::::::::::::::::::::::::::::: ::::::::. ::::::::::::::::::::::::::
8:.
::::::::::::::::::::::::::::::::::::::
.....................~:i:i:i i li:;:::i::!ii::ii i
3 4 5 6
135 KM 130 KM I 90 KM J lE K M - -
':::::::.:.... ~ ~..,
........................... !:::!: ~ ::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::: .:
. . . . . c~ i ~ (2) LOWERLAGUNILLA! ~ i i i i i
_: (3) LA ROSA
(41 ICOTEA
J~ : :SEAMROGIFORMIS ZONE::,
b ' . ~ : ' : + ~:':"'~:':~::':'~:'>:':~':':':':':':':':':':': Z,:c,
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: ~ ........
iiiii/}~illi~i!~,~!.....................iiiif:i........ LOCATION MAP
NARACAIBO
::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::: ::::::::::::::::::::::::::
/ V EN EZ U EL A
V. USMENS
S
I ZONE /
mBOGOIA
o 200 300 KM
I | I I
7 8 q
120 KM J 1 O0 KM I
DIEDERIXI Z O N E
GLOBOROIAEIA
MENARDII ZONE
GLOBOROIAUA
MAYERI ZONE
G. M A G N A C L A V A T A Z O N E
" GLOBOROIALIA
FOH$1 ROBUSTA ZONE
GLOBOROTAUA
EOHSI EOBATA ZONE
M VANDERHAMMENI ZONE
.................................
i 6EOBOROTALIA
L. FOHSI FOH$1 ZONE
POZON ................................
GLOBOROTALIA
i
,~ P. H I N I H U S Z O N E FOHSI BARJSANENSISZONE
..................................
GLOBIGERINAEELLA
0 -- INSUETA ZONE
........................
o EATAPSYDRAX
z ~.o J. SEAMROGtFORMIS ZONE SIAtNFORTHfl ZONE
"( ~ ......................
CATAPSYORAXDISSIMILIS ZONE
GEOBOROEALIA KUGLERIZONE
GLOBIGERINA
C. DOROGENSIS Z O N E
(IPEROENSIS CIPEEOENSISZONE
LEGEND
MARINE
SECTION II
1 2 3
I 200 KM I 400 KM I
P. DIEDERIXIZONE
ZONE
::::::::::::::::::::::::::::::::::::::::::::::::: z
METRESVERTISCALEFEE1
CAL ~...~................~..~::::::::::::::::::::::::::::::::::::::::::::::::
'~
0 0 :::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::
1000
2000 :~:~:~:~:~:~:~:~:!:~:~:~:~:~:i
"'"'""'"'"~" :~:~::-, ~ 1
3000
1000
4000
1.500 5000
4 5 6
BlO O N I A
/
/
/ LOCATION MAP
/ o
~ CARIBBEAN SEA
COLOMBIA
//
GLOBOROTALIA
N~NARDfl ZONE
GLOBOROTALIA
NAYERI ZONE
b GLOBOROTAUA
0 FOHSI ROBUSTA ZONE
..............................
~r GLOBIGERINATELLA m ~ ~ ~ ~ ~ ~ --
o - - ~ INSUETA ZONE
o ~ ~ ...........................
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GLOBIGERINA
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1 2 3
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P. DIEDERIXI Z O N E
G. M A G N A C L A V A T A ZONE
M. VANOERHAMHENIZONE ~:;::~.:.
..................
:.:.:.:.:.:.:.. ................................. Z ,~
::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::
VERTICAL SCALE ""::':-:':':'::':-:':':':'::-:-:':-:':-:':-..-:::
MEFRE FEEl
0 -- 0 ::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::: <
p MiN,i,iC~sz6NEili:i:i:i:i:i:i:i:i:i:i:i:
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2000
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3000
1 ooo ...............:::~:~:~:~!~;~:.{{~:::.::i:::.
4000
1500
5000
sedimentation is not excluded, although the rather sharp and distinct floral change
at the base of the Retibrevitricolpites triangulatus Zone still suggests the presence
of a hiatus. It increases southwards since, in Concepci6n, the Foveotricolpites per-
foratus Zone is missing, while in the North Central Lake area the Ctenolophondites
ilsamae Zone is also missing, and it reaches maximum values along the northern
Andean foothills. In Rio Mullapas and Rio Perdido the Proteacidites dehaani
Zone is found underlying the Eocene, and the whole Paleocene appears to be
missing.
In a southwesterly direction the magnitude of the hiatus rapidly decreases
again and in Quebrada La Mora the Foveotricolpites petforatus Zone is found
again. Further south and in the Tarra area the top of the Paleocene interval co-
incides with the base of the Mirador Sandstone, and it is likely on geological
grounds that a minor unconformity is present at this level (ScHAuB, 1948, p.225),
coinciding with the base of the Retibrevitricolpites triangulatus Zone. In Colombia
sampling gaps around this level preclude any definite statements on this problem.
The Eocene sedimentation cycle starts with the Retibrevitricolpites triangula-
tus Zone. (As mentioned in the previous section of" this paper the lowermost part
of this zone may be of Paleocene age.) Sediments assigned to this zone are thickly
developed in the northeast and in Colombia, but they are absent along the northern
Andean foothills between Rio de Oro and Rio Mullapas. The next higher Psilatri-
colpor#es crassus Zone is absent between Rio Perdido and Rio de Oro and in the
Colombian sections. The overlying Psilatricolpites operculatus Zone is generally
rather thin and its absence in sections with reduced sedimentation such as Rio
Mullapas and Quebrada La Victoria may be due to sampling gaps. In contrast
to the rather restricted areal distribution of the older Eocene zones, it is clear that
the overlying Retitricolporites guianensis and especially the Verrucatosporites
usmensis Zones have a much wider distribution. The former is thickly developed
in the Rio Mullapas and Rio Lebrija sections, while the latter is found in more or
less uniform thickness m all the northern Andean foothills sections as well as in
T~ichira. The absence of one or both zones in Rio Mullapas, North Central Lake
and Concepci6n is due to truncation by the well-known post-Eocene unconformity.
From these observations it may be deduced that during the Eocene progres-
sively younger sediments transgressed over an eroded Paleocene surface, first
localized in the deeper parts of the sedimentary basin, but rapidly transgressing
over virtually the whole area of investigation in the upper part of the Retitrico#
porites guianensis Zone and especially during the time of deposition of the Ver-
rucatosporites usmensis Zone. This last transgressive phase coincides with the
widespread deposition of the marine Pauji in the east, which sharply contrasts
with the transitional environment prevalent in the older Eocene.
It must be noted that in the southwestern part this transgressive younger
Eocene rests unconformably on Lower Eocene sediments of Retibrevitricolpites
triangulatus-Psilatricolporites crassus Zone age, indicating a minor orogenic phase,
BOTANICAL RESULTS
FILICALES
PARKERIACEAE
The source areas of this spore species are the alluvial plain and coastal
swamps where the parent plant grows in the shape of a small aquatic fern in
shallow water, bordering lakes and river banks. This environment is characterized
by rapid local facies changes and this is clearly reflected in the pronounced quan-
titative fluctuations in fossil coastal plain sediments. In marine sediments the
spores are much rarer. The taxonomic identification is thus well supported by the
distribution pattern of the fossil spore.
The frequency of Ceratopteris spores in the Neogene of all three areas
investigated and their absence in older Palaeogene and Cretaceous deposits, shows
that the present-day pantropical distribution of this freshwater fern may date only
from the mid-Tertiary. This is, however, somewhat in contrast to the isolated
systematic position and specialized morphology of the genus, which is assumed
to be of great antiquity. Its origin may have been local, but as long as no ancestral
spore forms have been found this problem remains unsolved.
SCHIZAEACEAE
This spore type is found in most of the species of Mohria and a few of
ANGIOSPERMAE
ARECACEAE(PALMAE)
BETULACEAE
PROTEACEAE
The presence of this pollen type in Senonian sediments of Nigeria and the
Caribbean would indicate a formerly more extended area of certain genera of
Proteaceae nowadays mainly restricted to the Southern Hemisphere (Guevina and
Lomatia in Chili, Leucospermum in South Africa, Lomatia, Cardwellia and Steno-
OLACACEAE
The earliest record of this highly characteristic pollen type dates from the
Middle Eocene in the Caribbean, where it remains present up to Recent. In Nigeria
the first appearance is in the Upper Eocene and it is remarkably restricted in its
range there, disappearing again in the Lower Miocene.
In Borneo it is absent in Cretaceous-Paleocene sediments, but its base is
not accurately known. In the Neogene it is fairly common, however. RAMANUJAM
(1966) has recorded similar pollen types from the Miocene of south India. Since
a number of genera may be represented, it is difficult to be positive about the
phyto-geographical significance of these facts.
It is possible that Caesalpinia bonduc, which prefers a coastal habitat, has
produced a large proportion of the fossil pollen of this type, but in the Caribbean
area Caesalpinia coriaria, which is common in the dry thorn forests along the
northeastern coast of South America, may also have contributed. Further detailed
studies of this group of pollen types will no doubt result in interesting conclusions.
uyACl~AE--Ctenolophoniideae
Ctenolophon
MALPIGHIACEAE
The fossil record for this characteristic pollen type, which occurs in several
genera of Malpighiaceae, dates from the Middle Eocene of the Caribbean area.
In Nigeria it appears slightly later and remains much more scarce than in the
Caribbean area. This is in agreement with present-day distribution of the family,
which is best represented in South America, in which continent its origin may have
lain.
Fruits of Malpighiaceae recorded from the Tertiary of Europe have so far
not been accompanied by fossil pollen. The less characteristic pollen of the Mal-
pighiaceae growing in Borneo today, has not yet been found fossil.
EUPHORBIACEAE
The highly characteristic pollen grains of Alchornea are first found in the
lower part of the Middle Eocene of the Caribbean area. In Nigeria they appear
somewhat later, in the upper part of the Middle Eocene. In Borneo their first
origin cannot yet be precisely given. It would appear that the present-day pan-
tropical distribution of the genus was achieved within a comparatively short inter-
val of time in the Eocene.
This pollen type appears at the base of the Eocene in the Caribbean area
and in Nigeria. It is most likely that the genus Amanoa, which occurs both in
BOMBACACEAE
The first occurrence of the genus Bombax can be dated as Paleocene in the
Carribbean area. No ancestral types are known, however, and the rather sharp
lower limit of occurrence suggests immigration from elsewhere. In Nigeria similar,
but not identical, pollen types are known from the Paleocene-Eocene transition
onwards. In the Caribbean area the Bombax ceiba type becomes rare during the
Eocene and has given rise to a host of related types which cannot be discussed
here because of their limited stratigraphical interest.
The very characteristic Catostemma pollen grains are not found below the
Upper Eocene in the Caribbean area. They show strong local dominance especially
in the Oligocene and Lower Miocene, which may be taken as an indicator of the
presence of rain forest. The origin of this pollen type is not known, but related
tricolporate pollen grains referable to the genus Aguiaria occur already in the
Lower Eocene.
CLUSIACEAE (GUTTIFERAE)
LYTHRACEAE
The typical pollen grains of the genus Cuphea, which is restricted to America,
first appear in the palynological record of the Caribbean area in the Middle Mio-
cene, but no ancestral forms are known.
In the Caribbean area Crenea pollen is first found in the Upper Eocene and
is especially common in the Lower Miocene, associated with dominant Rhizophora
pollen in coastal sediments. In Nigeria it first appears at the base of the Miocene
and is fairly abundant during the Lower Miocene, also in a coastal environment,
only to disappear from the stratigraphical record in the Middle Miocene. In the
Caribbean area a decrease sets in at approximately the same time, but Crenea
pollen remains present in small numbers up to the present day.
The palynological data thus provide a fairly complete record of the history
of the genus Crenea. Its origin must be sought in the Eocene of South America
and it apparently succeeded in crossing the Atlantic Ocean, probably during the
Oligocene. It then flourished for a short time on the western African coast in the
same environment as in South America. The reasons for its disappearance from
Africa and its almost simultaneous decrease in abundance in the Caribbean area
are unknown. As in the case of Symphonia, the coastal habitat of Crenea no doubt
was a principal factor which significantly increased the chances of dispersal.
SONNERATIACEAE
Sonneratia
RHIZOPHORACEAE
Included in this type are the pollen grains of the genera Rhizophora, Bru-
guiera and Ceriops. Although differences in pollen type are known to exist between
the genera and constituent species, it has proved impracticable to try to separate
them on a routine basis. The following account therefore relates mainly to the
history of the tribe Rhizophoreae, to which the genera mentioned belong and
which are all mangroves.
As far as known the Rhizophora pollen type is unique and cannot be confused
with pollen from other taxa (cf. also MULLER, 1964). The fossil record in Borneo
shows that the Rhizophora type is absent in Cretaceous and Paleocene sediments.
It probably occurs first in Eocene sediments, but due to poor preservation this
cannot yet be stated with certainty. In the Oligocene it is definitely present, but
in small quantities only; it gradually increases in abundance to become in the
Miocene-Pliocene the dominant element in the coastal microflora.
In the Caribbean area it is definitely known for the first time in Upper
Eocene sediments, being absent in older strata. Again it is only in the Miocene
that Rhizophora reaches the high percentages so characteristic of the Neogene and
Recent sediments of coastal or marine origin.
In Nigeria, in contrast, it is definitely absent from pre-Miocene sediments
and starts occurring rather suddenly in high percentages in the lowermost Miocene.
If this fossil evidence is combined with the present-day distribution pattern
of the tribe Rhizophoreae as discussed recently by VAN SVEENIS (1962a), the fol-
lowing tentative history can be reconstructed:
(1) Origin of the tribe Rhizophoreae in the Eocene of southeastern Asia,
since today the largest number of genera and species occur there, as well as inland
relatives (Carallia, Anisophyllea).
(2) Extension of the range of a few species of Rhizophora eastwards across
the Pacific Ocean to tropical South America, reaching the Caribbean area via the
gap in the Panama isthmus, most probably during the Eocene, followed by develop-
ment of local species.
(3) Crossing of the Atlantic Ocean in the Early Miocene and settling of the
American species of Rhizophora on the west coast of Africa.
(4) Extension of the range of the Indo-Malesian species of Rhizophora
westwards to east Africa at an as yet undetermined time.
ACANTHACEAE
Although some of the genera which produce this pollen type have a pan-
tropical distribution, the occurrences in the Neogene appear to be restricted to
the Caribbean area and Nigeria. It is possible that the extension of the range to
the Indo-Malesian area took place relatively late in the Mio-Pliocene.
The genera which produce this pollen type are restricted at present to the
American tropics and accordingly the fossil pollen type is known only from the
Caribbean area.
From the fact that acanthaceous pollen types are only known from the
Neogene it might be inferred that this family is of relatively late origin.
The following three Asteraceae pollen types have been distinguished: (l)
Tubul([torae type (Echitricolporites spinosus); (2) Liguliflorae type (Fenestrites
spinosus); and (3) Ambrosia type (Echitricolporites mcneillyi).
Future detailed studies on well-preserved fossil material will no doubt lead
to a further subdivision of these type groups.
The Tubuliflorae type appears to be the oldest and the most widespread of
the three Asteraceae pollen types. It has not yet been found with certainty in
pre-Miocene sediments.
In all three areas investigated the very gradual increase in abundance appears
to be contemporaneous, but the high percentages reached in the Mio-Pliocene of
the Caribbean area have not been found so far in Nigeria or Borneo.
The Liguliflorae type is much rarer and has only been regularly observed in
the Caribbean area, where it appears slightly later than the Tubuliflorae type.
The Ambrosia type, only known from the Caribbean area, appears still later.
Asteraceae are widespread in the present-day tropics, but are more common
in open vegetation types, such as the savannah or higher montane vegetation than
in the closed lowland rain forest. This largely explains the considerable quantities
of Asteraceae pollen observed in the Caribbean area Mio-Pliocene in comparison
with the relative scarcity in an area such as northwestern Borneo.
Earlier views (KUvL et al., 1955) that, on the basis of the palynological
record, Asteraceae emerge as one of the youngest developments within the angio-
sperms are thus confirmed. The place of origin is still obscure, however, and may
very well have been situated outside the tropical belt, the time of origin presumably
being mid-Tertiary.
The relatively quick and simultaneous extension of the range of Asteraceae,
especially of the Tubuliflorae type, as shown by the palynological data, probably
reflects the efficient seed dispersal which characterizes the family.
SUMMARY OF BOTANICALRESULTS
PLATE I
i i i Iii
PLATE II
f~
proximal face psilate, surrounded by a circular ridge, which makes contact with
the striate ridge pattern at the points of the laesura. Remainder of wall coarsely
striate; striae 1-2/~ high, 2-3/z wide; grooves 1~-3 ~t wide. Exine 1~-2~
~ it thick.
Dimensions." 77 132/z (equatorial diameter).
Variability: There is some variability in size and sculpture; thicker striae
may occur in smaller number or much wider grooves. Also the proximal side of
the grain may be locally perforated.
Distribution: Regularly present from the base of the Magnastriatites howardi
Zone upwards, in all three areas. Its first appearance is remarkably sudden and,
as far as can be judged simultaneous. No ancestral forms are known.
Taxonomic affinities." Virtually identical with the spores of the tropical-
subtropical fresh-water fern genus Ceratopteris (species seen: C. thalictroides,
Plate IV, 1, and C. cornuta).
Comments: The spores described by VAN DER HAMMEN (1956d) under the
genus Verrumonoletes clearly fall within the circumscription of Verrucatosporites
and this name has priority.
P L A T E III
Derivatio nominis." The genus has been named in memory of the late Dr.
T. F. Grimsdale, who launched the idea of applying palynology in oil exploration
in 1937.
Type species." Grimsdalea magnaclavata nov. sp.
Diagnosis: Spherical, aperture indistinct or absent, wall thin, intectate,
sculpture of two types, finely scabrate and coarsely clavate.
PLATE IV
Literature." Monoporites annulatus VAN DER HAMMEN, 1954, p.90, pl.6, fig.4.
Monoporites unipertusus VAN DER HAMMEN, 1956b, p.82, pl.5,
fig.10.
Description." Single grain, radially symmetrical, anisopolar, almost spherical.
Single aperture small, circular, penetrating entire wall, costate; costa 4~-/~ wide,
slightly protruding. Endexine < /~ thick; columellae indistinct <: /~ long and
thick; tectum < # thick, psilate-very finely perforate or scabrate.
Dimensions." 38-43/~.
Variability: Considerable variability in size of grain and pore exists, but
the wall is always rather thin. The surface of the wall is mostly smooth but a finely
scabrate surface occurs occasionally. Since it is mostly unknown whether this
condition is due to corrosion or not, this sculpture type is included.
Distribution: In the Caribbean area and in Nigeria occurring regularly from
the base of the Monoporites annulatus Zone upwards, although with rather pro-
nounced fluctuations in numbers and generally more abundant in the Neogene
than in the Palaeogene. Rare occurrences below the base of the Monoporites
annulatus Zone are known. In Borneo base not yet determined, but absent in
Paleocene and Upper Cretaceous. The significance of the fluctuations of grass
pollen has been discussed in the section "Botanical results".
PLATE V
PLATE VI
Diagnosis: Pollen grains with two ectexinous colpi combined with two
endexinous pores.
Derivatio nominis: Named in honour of Dr. Th. van der Hammen in recog-
nition of his contributions to South American palynology.
Holotype: Slide TC-155, well Hervidero-l, 2770 ft., Venezuela.
Description: Single grain, 90 rotated bilaterally symmetrical, isopolar,
spherical; dicolporate; colpi very long, ectexinous, bordered by 3-4 strips, 4-5/z
wide, perpendicularly orientated, straight borders and pointed or rounded ends;
pori endexinous, transversely elongated, 4/z wide, 13 # long, constricted in the
middle, costate; costae strongly protruding, 1-2/~ thick. Endexine < 1 /z thick;
columellae 1 # long, /z thick, tectum 2/z thick, foveolate, lumina I~-2 # wide,
muri 2-3/z thick.
Dimensions: 40-53/~.
Variability: In size and coarseness of sculpture.
Distribution: Only known from the Caribbean area where it occurs from
the base of the Retitriletes vanraadshooveni Zone upwards.
Taxonomic affinities: Close resemblance exists with Sanchezia klugii and
Trichanthera gigantea (Plate VI, 7; Plate VII, 1). The latter grain, however, has
a considerably larger size (105 #!). Other multimarginate species of Acanthaceae
available are non-rotated symmetrical (Asteracantha longiJblia, Asystasia
vogeliana, M imulopsis solmsii).
Derivatio nominis: The genus has been named in memory of the late Prof.
Dr. F. Florschfitz, the founder of palynology in The Netherlands.
Diagnosis: Subprolate, sometimes trilobate and with meridional ridges.
Pores equatorial, circular, distinct. Wall tectate, columellae generally indistinct,
sometimes fused. Tectum continuous, smooth or broken up into separate verrucae
and differentiated in polar and/or meridional direction.
Type species: F/orschuetzia tri/obata nov. sp.
Comments: The genus differs from Verrutrico/porites VAN DER HAMMEN in
the absence of colpi, the indistinctly columellate structure and the absence of a
supratectate verrucate sculpture. The diagnosis has been taken wide enough to
include the fossil sonneratioid pollen types, including the presumed ancestral one.
Some confusion may, however, be possible with a "sporotype" described in 1954
by COOKSON and PIKE and named Santa/umidites, the description of which has
subsequently been emended and accepted as a valid form genus by POTONI~(1960).
In both Cookson and Pike's and Potoni6's diagnosis it is clearly stated that the
maximum thickness of the wall is around the pores, as is the case in Santa/urn
pollen. No confusion with Florschuetzia appears likely, therefore, and the genus
would not have to be mentioned here, but for the fact noted before (MULLER,
1964) that the photomicrographs accompanying Cookson and Pike's paper strongly
suggest that fossil pollen of Sonneratia caseo/aris has been included in S a n t a & m #
dites (see especially their fig. 71, pl. 2, which is strikingly similar to F/orschuetzia
levipoli). This suspicion is strengthened by Cookson and Pike's discussion of the
affinities of Santalumidites cainozoicus in which the authors mention puzzling
differences with recent Santalum pollen and a large degree of variability as well.
If this would indeed prove to be the case, there would have been no choice
left, other than correcting Cookson and Pike's description of the type and accepting
P L A T E VII
PLATE VIII
PLATE IX
~:IIm!i~ ~ ~ i~ ~ ~:::~mm
rll~ 11mmmm
i~~ ~i~i~i~~ ~ i~~ ~i!i~i!i!ii!~
~::~i~!i,,,,~,~,~i~i~!~,~:~
~ , i~ ~,~,~,ii~i~,~ii~,
~?~ii~i~,~,j~i~iiii~ii:~:,~S~i~!~!~J~
~,~: ~ ~
Literature: Echitriporites trianguliformis VAN HO~KFN-KLINKEN~ERG,1964,
p.218, pl.47, fig.7.
Echitriporites trianguliformis BELSK, BOLTENHAGENet POTON~k,
1965, p.77, pl.13, fig. 22, 23.
Description: Single grain, radially symmetrical, isopolar, oblate; outline in
polar view semi-angular. Triporate, pores ectexinous and endexinous, circular,
2 # in diameter, costate, often slightly protruding. Endexine 4 1 /z , intectate-
echinate, echinae 1-2 ,u long, ~: 1 /z wide at base, conical or with rounded tops,
2/z apart.
Dimensions: 21-30 #.
Variability: Considerable variation exists in size and shape of the grain and
in size and density of the spines. The more triangular grains with fewer and smaller
spines are more common in the Upper Cretaceous of northern South America,
whereas the more rounded grains with slightly more and larger spines are more
common in the Eocene. However, since many transitional specimens occur, no
specific distinction was possible.
Distribution: Encountered in the Proxapertites operculatus Zone in all three
areas, but rare in Borneo and soon disappearing. In the Caribbean area and
Nigeria the species remains an important component of the Eocene microfloras
which has great practical value for local correlations. In both areas it disappears
from the record around the top of the Verrucatosporites usmensis Zone.
Taxonomic affinities: Despite intensive search, no positive identification has
been possible yet. This, and its gradual disappearance at the top of the Eocene
suggest that the taxon producing the pollen grains is probably extinct. A superficial
resemblance is shown by some Proteaceae (Embothrium, Garnieria, Persoonia,
Telopea). Only Telopea oreades and Embothrium mucronatum were available for
comparison but these species showed finely columellate-tectate pollen grains.
PLATE X
PLATE XI
1. The~pesiapopulnea COgREA,Recent.
2. Buttinia andreevi BOLTENHAGEN,Colombia, upper focus.
3. Buttiniaandreevi BOLTENHAGEN,Colombia, lower focus.
Magnification 1,000.
Remarks: Since Van der Hammen has invalidly indicated a recent pollen
grain as type specimen, his genus is hereby legalized by selecting as lectogenotype
the fossil species as described below.
Literature: Fenestrites spinosus VAN DER HAMMEN, 1956b, p.97, pl.12, fig.38.
Description: Single grain, radially symmetrical, isopolar, spherical; outline
In polar view almost hexangular. Colpi and pori indistinct, probably tricolporate.
Exine differentiated into a pattern of intectate lacunae (fenestrae), 8-11 /~ wide
and tectate-columellate cristae. Columellae 1 # long, /z thick; cristae 1-1 #
wide, 1/t high, bearing single rows of spines, 2 # long, 1 # wide at base.
Dimensions: 30-38 # (including cristae, excluding spines).
Variability: In size and coarseness of ornamentation. Sometimes undulating
cristae have been observed.
Distribution: Restricted to the middle and upper part of the Echitricolporites
spinosus Zone. More frequently observed in the Caribbean area than elsewhere.
Taxonomic affinities: This species is typical for the liguliflorae pollen type
of the Asteraceae (Compositae), which is produced by a large number of genera.
Of the many species which produce this pollen type, the following only will be men-
tioned as coming close to the fossil grains: Elephantopus angustifolia, Rolandia
fruticosa (Plate XII, 3), Vernonia canescens, Vernonia remotiflora (Plate XII, 2).
PLATE XII
1. Fenestrites spinosus VAN DER HAMMEN,Venezuela.
2. Vernonia remotiflora L. C. RICHARD,Recent.
3. Rolandia fruticosa (LINNAEUS)KUNTZE,Recent.
4. Striatricolpites catatumbus GONZALEZ,Colombia.
5. Crudia amazonica SPRUCE, upper focus, Recent.
6. Crudia amazonica SPRUCE, lower focus, Recent.
7. Foveotricolpites perforatus VAN DERHAMMENet GARCIA,Venezuela, upper focus.
8. Foveotricolpites perforatus VANDERHAMMENet GARCIA,Venezuela, lower focus.
9. Foveotricolpites perforatus VANDERHAMMENet GARCIA,Venezuela.
10. Perfotricolpites digitatus GONZALEZ,Venezuela.
Magnification 1,000.
4 !~-T~~i~ 5 ii ~' 6
i ~i~,iiiii!ii~i)!!~!!~i~i~~i~~i~ ~!!ii!iilJii!iif
~ l O
PLATE XIII
1. Scaevola plumieri VAI4L,Recent.
2. Merremia glabra HALUER(ill.), Recent.
3. Stephanoeolpites eostatus VAN DER HAMMEN, Venezuela.
4. Tabernaemontana attenuata URBAN,upper focus, Recent.
5. Tabernaemontana attenuata URBAN,lower focus, Recent.
6. Gemmastephanocolpites gemmatus VAN DER HAMMENet GARCIA,polar view, Venezuela.
7. Gemmastephanocolpites gemmatus VAN DER HAMMENet GARCIA,equatorial view, Venezuela.
Magnification 1,000.
,~,~iii,~ ~
x
Dimensions: 44--64/z.
Variability: A well characterized species with some variability in size and
coarseness of wall structure.
Distribution: Restricted to the Foveotricolpites perforatus Zone of the Carib-
bean area.
Taxonomic affinities: Unknown.
PLATE XIV
1. Retistephanocolpites williamsi nov. sp., Nigeria, holotype, upper focus.
2. Retistephanocolpites williamsi nov. sp., Nigeria, holotype, lower focus.
3. Ctenolophon parvifolius OLIVER,upper focus, Recent.
4. Ctenolophon parvifolius OUVER,lower focus, Recent.
5. Ctenolophonidites costatus VAN HOEKEN-KLINKENBER~,Nigeria, upper focus.
6. Ctenolophonidites costatus VAN HOEKEN-KUNKENaERG,Nigeria, lower focus.
7. Ctenolophonidites lisamae VAN DER HAMMENet GARC~A,Venezuela, polar view.
8. Ctenolophonidites lisamae VAN DER HAMMENet GARCIA,Venezuela, equatorial view.
Magnification 1,000.
~ i ~i~!ii~
~ _.
~ ~:~i~ ~~:i~~!~
1!
Description: Single grain, radially symmetrical, isopolar, spherical-sub-
oblate; outline in polar view stellate-circular. 5-colpate, colpi 14 .u long, ectexinous,
slightly costate equatorially. Ectexine locally thickened into a pattern of radial
costae, joining in pairs around the terminations of the colpi, not touching near
poles; ridges 12~--2bt high, 2-3 ,u wide. Columellae and tectum not differentiated;
wall finely scabrate, 3-4 # thick at equator, apparently densely perforated in all
directions, creating a spongy effect which diffuses the light considerably.
Dimensions: 17-31 #.
Variability: Some variability is present in the way in which the ridges unite
with each other.
Comments: In view of the close relationship which exists between this species
and Ctenolophonidites costatus, we have transferred it to the latter genus. The
identity of our specimens has been confirmed by Van der Hammen.
Distribution: Virtually restricted to the Ctenolophonidites lisamae and
Foveotricolpites perforatus Zones in the Caribbean area.
Taxonomic affinities: Has in all probability been derived from an extinct
species of Ctenolophon (Ctenolophonaceae).
Genus Psilatricolporites (VAN DER HAMMEN, 1956) ex VAN DER HAMMENet WYM-
STRA, 1964
PLATE XV
1. Ctenolophon engleri MILDBREAD, upper focus, Recent.
2. Ctenolophon engleri MILDBREAD, lower focus, Recent.
3. Psilatricolporites operculatus VAN DER HAMMENet WYMSTRA, Surinam.
4. Alchornea cordifolia MUELLER-ARG., Recent.
5. Alckornea obovata PAX et HOFFMANN, Recent.
6. Zonocostites ramonae nov. gen., nov. sp., Borneo, holotype.
7. Zonocostites ramonae nov. gen., nov. sp,, Borneo, holotype.
8. Rhizopkora rnucronata LAMARCK,upper focus, Recent.
9. Rhizophora mucronata LAMARCK,lower focus, Recent.
Magnification 1,000.
. . . . ~ ~i~iii~ii~,~'~' i~,~!,~i~!~ii~!!~ii,'
d d m L ~ ,, ~. ~., ~ . ~. ~ . .I F . . . ~
~ i ~ ~~i~i~ii~!~i!~!i~,~,,,,,,,,,,i,,i, ,~ !~iii~i~iii~iii~,'~iii,~'i~i~ili~,i'~'~!i~,,i~
PLATE XVI
1. Psilatricolporites crassus VAN DER HAMMENet WYMSTRA,Nigeria.
2. Psilatricolporites crassus VAN DER HAMMENet WYMSTRA,Venezuela.
3. Hura crepitans LINNAEUS, upper focus, Recent.
4. Hura crepitans LINNAEUS, lower focus, Recent.
5. Verrutricolporites rotundiporis VAN DER HAMMENet WYMSTRA, Trinidad.
6. Verrutricolporites rotundiporis VAN DER HAMMENet WYMSTRA,Surinam.
7. Verrutricolporites rotundiporis VAN D~R HAMMENet WYMSTRA, Nigeria.
8. Crenea maritima AUBLET, Recent.
9. Crenea maritima AUBLET, upper focus, Recent.
10. Crenea maritima AUaLE'r, lower focus, Recent.
11. Echitricolporites spinosus VAN DER HAMMEN, upper focus, Venezuela.
12. Echitricolporites spinosus VAN DER HAMMEN, lower focus, Venezuela.
13. Riencourtia glomerata CASS, upper focus, Recent.
14. Riencourtia glomerata CASS, lower focus, Recent.
Magnification 1,000.
Genus Retitricolporites (VAN DER HAMMEN, 1956) ex VAN DER HAMMEN et WYM-
STRA, 1964
P L A T E XVII
~1 l~+i+!+?
+++++++ + m, +.
Literature: Retitricolporites guianensis VAN DER HAMMENet WYMSTRA,1964,
p.235, pl.IH, fig.l-2.
Description: Single grain, radially symmetrical, isopolar, prolate. Tricol-
porate; colpi ectexinous, long and intruding with straight, marginate borders and
pointed ends; pores endexinous, circular, 1-2 # wide, rather indistinct. Endexine
/z thick; columellae /z thick and long, regularly distributed and rather closely
spaced; tectum # thick, reticulate; muri /~ thick and high, multicolumellhte,
but not always distinctly visible, lumina of angular shape, slightly elongated,
3-4/z wide, diminishing in size towards colpi.
Dimensions: 28-47/z.
Variability: Size and shape of this species are rather variable, the smaller
specimens showing a finer ornamentation. The species can, however, be easily
recognized by its loose-meshed reticulate sculpture with angular shape of the
lumina and thin, low muri. In well preserved specimens the regularly distributed
columellae are characteristic.
Comments: Inspection of type material has shown the identity of the speci-
mens described here with Retitricolporites guianensis. Our description, however,
differs slightly from the one given by VAN DER HAMMEN and WYMSTRA(1964).
Distribution: Only known from the Caribbean area where it occurs from
the base of the Retitricolporites guianensis Zone upwards.
Taxonomic affinities: A certain resemblance exists with pollen of Firrniania
colorata and Hildegardia barteri (Sterculiaceae), but a striking difference is the
finely reticulate appearance of the fairly broad muff in these types. To a lesser
degree this is the difference also with pollen of Glossosternon bruguieri (Sterculia-
ceae). The pollen of Pterocyrnbiurn beccarii has finely reticulate muri and in
addition a thicker wall. Pollen of Sterculia rnexicana has a thicker wall and is finer
reticulate on the poles. Trichospermurn pollen (Tiliaceae) has distinctly equatorially
elongated pores, although the wall structure is strikingly similar.
In view of this situation it is not yet possible to come to a firm decision
which taxon has produced Retitricolporites guianensis pollen. However, Ster-
culiaceae and Tiliaceae are the only families in which this pollen type has been
found so far.
PLATE XVIII
~ ~ili~i~ii~i ~ ~i
i
which is not mentioned by VAN DER HAMMENand GARCIA(1966), as an additional
typical feature.
Distribution: In the Caribbean area restricted to the Ctenolophonidites
lisamae and the Foveotricolpitesperforatus Zones. Absent from Borneo and Nigeria.
Taxonomic affinities: Closest resemblance is with certain species of the genus
Bombax (Bombacaceae). Especially close are Bombax ceiba (Plate XVII, II, 12),
B. rhodognaphalon, and B. pubescens, which all show the multicolumellate structure
in varying degrees. Bombax mexicana differs in the pronounced triangular shape.
No other genus of Bombacaceae shows this pollen type and identification with
a, possibly extinct species of Bombax appears well founded.
Derivatio nominis: The genus has been named in honour of Prof. J. Dufour,
who successfully stimulated the early palynological investigations in Venezuela.
Diagnosis: Oblate, 4-6 colporate, colpi ectexinous, slightly costate, reaching
approximately halfway the poles, endexinous pores indistinct, equatorially elon-
gated, wall rather densely columellate, tectate-densely perforate, perforations
evenly distributed, lumina < /~ in width.
Type species: Jandufouria seamrogiformis nov. sp.
Comments: This genus accommodates fossil pollen of the Catostemma type.
It can be distinguished from Retistephanocolporites by its evenly and densely
perforated tectum.
Derivatio nominis: Name derived from the resemblance in shape to the leaf
of the Irish shamrock (Trifolium repens).
Holotype: Slide TC-167, well Aurora-l, 651 ft., British Guyana.
Description: Single grain, radially symmetrical, isopolar, oblate. 4-6 col-
porate; colpi ectexinous, 36/z long, reaching half-way the poles, slightly costate
with straight borders and pointed ends, costae 1~-2 # wide; endexinous apertures
equatorially elongated, faintly costate. Endexine 1 # thick; columellae -~-1 # long,
# thick; tectum -1 /~ thick, densely and evenly perforated.
Dimensions: 40-57/~.
Variability: There is some slight variability in size and in wall thickness, but
as a rule the species is highly characteristic because of its shape.
Distribution: Only known from the Caribbean area, where it occurs from
the base of the Verrucatosporites usmensis Zone upwards.
Taxonomic affinities: Close resemblance exists with the pollen of the genus
Catostemma (Bombacaceae), especially with C. altsonii (Plate XVIII, 7, 8), C.
sclerophyllum, and C. fragrans. C. commune has distinctly digitate columellae, a
feature which has not been observed in the fossil material so far. The pollen of
Aguiaria is similar, but tricolporate. The widespread occurrence of Catostemma
in the rain-forests of tropical South America supports the identification.
ACKNOWLEDGEMENTS
In the compilation of this paper the authors were able to draw on the accumu-
lated experience of a large number of colleagues, whose assistance and criticism
is gratefully acknowledged here. Thanks are also due to the Management of the
Bataafse Internationale Petroleum Maatschappij and the Shell-BP Petroleum
Development Company of Nigeria Ltd. for permission to publish this paper,
REFERENCES