Review of Palaeobotany and Palynology, 65 (I 990): 105-113 105

Elsevier SciencePublishers B.V., Amsterdam

Campanian to Paleocene spore and pollen assemblages of

Seymour Island, Antarctica

R o s e m a r y A. A s k i n
Department of Earth Sciences, University of California, Riverside, CA 92521 (U.S.A.)
(Received October 27, 1988; revised and accepted January 25, 1990)

ABSTRACT

Askin, R.A., 1990. Campanian to Paleocene spore and pollen assemblages of Seymour Island, Antarctica. Rev. Palaeobot.
Palynol., 65: 105-113.

Spore and pollen assemblages in Campanian to Paleocene nearshore marine to deltaic sediments of Seymour Island,
Antarctica, are characterizedby abundant podocarpaceousconifer pollen, diverseprovincialangiospermpollen and cryptogam
spores of low diversity. These assemblages resemble coeval assemblages from New Zealand, southeastern Australia and
southern South America. The palynoflora reflectsprimarily conifer-dominatedrainforest growing in cool to warm temperate
paleoclimates. The late Maastrichtian was a comparativelywarm interval, with a humid equable paleoclimate. The Antarctic
Peninsula cordillera probably supported altitudinally zoned plant associations and it is likelythat climatic fluctuationsduring
the Campanian to Paleoceneresulted in altitudinal shifts of these vegetation zones.

Introduction basal Danian beds (member/"unit" 10 of L6pez de

Spore and pollen assemblages from upper
Campanian to Paleocene sediments on Seymour
Island, Antarctica, provide a glimpse into the
Cretaceous/Tertiary land vegetation and climate of
the northern Antarctic Peninsula. The rich and
varied palynomorph assemblages also include
marine dinoflagellate cysts useful for biostratigra-
phy and interpretation of depositional environ-
ments. Diverse macro- and microfossil faunas and
floras associated with the palynomorphs provide
independent stratigraphic control.
Palynomorphs discussed here are from approxi-
mately 500 outcrop samples in measured sections
through the upper Campanian, Maastrichtian and
Paleocene L6pez de Bertodano and Sobral Forma-
tions (Fig. 1). Most of this succession was deposited
in low energy, nearshore marine paleoenviron-
ments. A regression, beginning near the end of the
Cretaceous, changed shallow shelf conditions in
most of the L6pez de Bertodano Formation to
inshore/estuarine depositional environments in the
Bertodano Formation). The overlying basal Sobral
Formation (members 1 and 2) reflects more marine
conditions, followed by shallowing to deltaic
conditions for the main part (members 3-5) of the
Sobral Formation (Macellari, 1988; Sadler, 1988;
Askin, 1988a). Nearly all the sediments ate
unconsolidated and fine-grained (sandy silts and
muds) and contain abundant, well-preserved paly-
nofloras, except members 3 and 4 of the Sobral,
which are predominantly sandy with sparse paly-
nomorphs.
The provenance of the sediments (and the spores
and pollen) was the northern Antarctic Peninsula,
a magmatic arc that underwent several periods of
plutonism, volcanism and uplift during the Creta-
ceous and Cenozoic (Elliot, 1988). This cordillera
supported plant associations in a range of habitats
from coastal to alpine.
The stratigraphic framework and dinoflagellate
cyst succession have been summarized for the
Campanian to Paleocene of Seymour Island
(Askin, 1988a). The spore and pollen assemblages

0034-6667/90/$03.50 © 1990-- ElsevierSciencePublishers B.V.

106 R.A. ASKIN

sE¥ ou, ,sLA,o 9.0e I m.p

I (southern p a r t ) /i!i! iiiliiiil!iiiiiiiiiiiiiii] ~[ Cross
=========================================== ~ ~ ~.
==================================:::-:~ o
0 I Z Km ~ /~i.li:i:?:i . . . . . . . . . . . i:i ........i:
" ~ J " ' " "' ........ '"'~ -- "1400- 5~

Ia t 1200.

:~ I Ioo-
i!0' 9
I ' ~ '~T, ~'~(~///1~///I ~ Iooo-

l~'~1 ~1 I ., ,"7 . . ~ ~ r.////l 5

F' t d l ,¢7 ,oo. o ,

.,--~ ............. ~l Sobral Fm. ¢ c " "~

South Shetland ! ~ ~ ~-vl~ u~=

Islands (~ ~ mud.stone J ' B e r t o d o n o Fm ~. o,~n
I • ~ ",,~ . . . . roc.es E ,.v,,- 3
• -,r~,,,.,4t3~O ~
~ " ~ Dykes P
V r E ~,~e.....-Vegn Is. ~ ' ~ OO-
~a64~sl ~'~/~eLI~..Seym our ^ 7 .................. ~ "-- 2
~'/" .,~,,~t~ } ( c . , , ~ v I s l and . . . o I
.,AI0,,~/V/-x~'/-"SO,now H#l ,s
~'/ / / ~ ond numbers -- 0
??- ~ oo~7 .oss ,~. ,

Fig. 1. Geologic map and stratigraphic column for southern part of Seymour Island. L6pez de Bertodano members after Macellari
0988) and Sobral members after Sadler 0988).

and their significance, including comments on the and stratigraphic ranges (within the late Campan-
Cretaceous/Tertiary floral turnover, are discussed ian to Danian) on Seymour Island.
below. The dominant species throughout the Campani-
an-Paleocene is Phyllocladidites mawsonii. This
Composition of palynofloras species, with other common conifer pollen (e.g.
Podocarpus and Dacrydium type) reflects conifer-
The Campanian-Paleocene spore and pollen dominated rainforest vegetation during the Cam-
flora of Seymour Island contains abundant podo- panian-Paleocene. Other conspicuous elements
carpaceous conifer pollen (average 40-65% of are pollen of proteaceous species and southern
terrestrial assemblage), other gymnospermous pol- beech (Nothofagus, including brassii, fusca and
len as minor elements (< 2%), varied and usually menziesii group species). The proteaceous species
subdominant angiosperm pollen (20-45%) and are common and varied, although not as diverse as
cryptogam spores (6-15%). Table I lists most of in Paleogene palynofloras of Australia. Species of
the spore and pollen species found in these Peninsulapollis a r e of presumed proteaceous affin-.
samples, with their probable botanical affinities ity, having similarities to the extant Beauprea
SPORE AND POLLEN ASSEMBLAGES OF SEYMOUR ISLAND
(Dettmann and Jarzen, 1988a). They are some of
the most common angiospermous pollen in the
Campanian-Maastrichtian.
These spores and pollen exhibit varying degrees
of provincialism (Askin, 1989). Some are endemic
to the James Ross Island basin or to Antarctica,
while others have a Weddellian Province distribu-
tion (southernmost South America, West Antarc-
tica, southeastern Australia and New Zealand), or
a wider southern (Austral) distribution. Cosmo-
politan taxa were also found.
The Campanian to Paleocene palynoflora of the
Weddellian Province reflects a podocarpaceous
conifer-dominated rainforest. P. mawsonii conifer
pollen is prevalent, along with secondary proteal-
eans, Nothofagus and other angiosperms and
moisture-loving (rainforest or riparian) crypto-
gams (e.g. Stover and Partridge, 1973; Archangel-
sky and Romero, 1974; Mildenhall, 1980;
Truswell, 1983; Raine, 1984, 1988; Dettmann,
1986; Dettmann and Thomson, 1987; Dettmann
and Jarzen, 1988a,b). Differences within this
province are also apparent (e.g. Dettmann and
Thomson, 1987).
Stratigraphic implications
As indicated in Table I, some spore and pollen
species have restricted stratigraphic ranges and are
potentially useful biostratigraphic markers for the
Antarctic-Subantarctic region. Weddellian Prov-
ince species allow broad correlations with south-
eastern Australian and New Zealand spore and
pollen zones. The lower and upper parts of the
Seymour Island late Campanian to Maastrichtian
succession (units 1-9 of L6pez de Bertodano
Formation) correspond respectively to the T. lilliei
and T. longus Zones of southeastern Australia
(Stover and Evans, 1973; Helby et al., 1987). Both
parts correlate with the PM2 Zone of South Island,
New Zealand (Raine, 1984). The Danian succes-
sion (L6pez de Bertodano 10 and Sobral 1-4)
corresponds to the Australian L. balmei and New
Zealand PM3 zones.
Species ranges are dissimilar in the different
parts of the Weddellian Province. This disparity in
ranges prevents precise correlation. For example,
on Seymour Island, first appearances of Tubuliflor-
107
idites lilliei and Tetracolporites verrucosus occur in
unit 1 (of L6pez de Bertodano Formation);
Triporopollenites sectilis and Grapnelispora
sp.cf.G, evansii (the latter is only 1/3 the size of the
Australian and New Zealand specimens) are in
unit 4; and Quadraplanus brossus is in unit 6. The
timing of some Australian first appearances (from
fig. 33 of Helby et al., 1987) differs. For example,
Triporopollenites sectilis appears before Tetracol-
porites verrucosus in Australia.
Dettmann (1986), in her comprehensive discus-
sion on the fern Lophosoria and its fossil record
(Cyatheacidites spores), warns of the dangers of
using terrestrial palynomorphs for regional corre-
lation. Plant migration that is dependent on
habitat availability may proceed over geologically
long periods of time. Dettmann (1986, 1989) and
Askin (1989) refer to austral species that have
migrated across the southern continents, produc-
ing a fossil record of disparate or "heterochro-
nous" first appearances along their dispersal
pathways. Some of these taxa, however, may still
be useful locally as biostratigraphic markers.
Campanian-Paleocene floral history
The preserved late Campanian to Paleocene
palynofloras represent different plant associations
intermixed (and probably reworked in part) by the
time they were deposited in the Seymour basin.
These composite assemblages contain mainly cool
temperate climate indicators, assuming climatic
preferences of modern analogues extend back to
the latest Cretaceous. Some warmer climate indica-
tors also occur, particularly in the late Maastri-
chtian. The vegetation was probably zoned by
altitude, with the "warmth-loving" (or more
equable) plant associations occupying
coastal/lowland areas and cool climate associa-
tions in the higher altitude, inland areas.
Many of the conifers, and species of the
Nothofagus fusca and N. menziesii groups, are
typical of cool temperate climates. The distinctive
conifer pollen Phyllocladidites mawsonii, invariably
the predominant species in the Campanian-Paleo-
cene, is identical to pollen of the extant Lagarostro-
bus (Dacrydium) franklinii (Cookson, 1953; Play-
ford and Dettmann, 1978). This is a rainforest tree
108 R.A. ASKIN

TABLE I

Pollen species in the late Campanian to Danian of Seymour Island.

Botanical affinity Spore and pollen species Range on Seymour Island

[presumed modern analogue]

Cryptogams
Hepaticae
Sphagnaceae (and other Musci) Species documented in Askin (in press)
Selaginellales Stereisporites spp. [Sphagnum] and Documented in Askin (in press)
Lycopodiales Laevigatosporites spp. [Filicales]
Filicales are most abundant spores
Salviniales
Gymnosperms
Cycadales/Ginkgoales Cycadopites spp.
Podocarpaceae Dacrycarpites australiensis Cookson &
Pike [Dacrycarpus]
Dacrydiumites spp. [Dacrydium]
Lygistepollenites florinii (Cookson &
Pike) Stover & Evans, L. balmei
(Cookson) Stover & Evans [Dacrydium]
Microcachryidites antarcticus Cookson
[cf. Microcachrys]
Phyllocladidites mawsonii Cookson ex late Campanian-Danian-
Couper [Lagarostrobus franklinil]
Phyllocladidites spp. [Phyllocladus]
Podocarpidites spp. [Podocarpus]
Podosporites spp. [Microcachrys/
Microstrobos]
Trichotomosulcites subgranulatus Couper
Araucariaceae Araucariacites australis Cookson
[Araucaria]
Ephedraceae Ephedra notensis Cookson late Maastrichtian-Danian-
Angiosperms
Liliaceae Liliacidites cf. kaitangataensis Couper late Campanian-Danian
Liliaeidites spp. Maastrichtian-Danian-
Aquifoliaceae llexpollenites sp. late Campanian-Danian
Bombacaceae Bombacacidites bombaxoides Couper late Maastrichtian
[Bombax]; Bombacacidites sp.
Casuarinaceae Haloragacidites harrisii (Couper) Harris Danian-
Ericales, ?Epacridaceae Ericipites sp. Danian-
Fagaceae Nothofagidites spp. [Nothofagus brassii, late Campanian-Danian-
fusca, menziesii groups]
Gunneraceae Tricolpites reticulatus Cookson late Campanian-Danian-
[Gunnera]
tauraceae 1 ?Dilwynites granulatus Harris, Danian
D. tuberculatus Harris [?Cinnamomum]
Loranthaceae Cranwellia striata (Couper) Srivastava late Campanian-late Maastrichtian
Cranwellia sp. late Maastrichtian
Myrtaceae Myrtaceidites spp. late Campanian-Danian-
Olacaceae Anacolosidites sectus Partridge latest Maastrichtian
[Anacolosa]
Proteaceae Proteacidites spp. late Campanian-Danian-
Propylipollis spp. late Campanian-Danian-
Cranwellipollis palisadus (Couper) late Campanian-M aastrichtian
Martin & Harris
Cranwellipollis subpalisadus (Couper) Maastrichtian-eady Danian
Martin & Harris
SPOREANDPOLLENASSEMBLAGESOF SEYMOURISLAND 109

TABLE I (continued)

Botanical affinity Spore and pollen species Range on Seymour Island

[presumed modem analogue]

Beaupreaidites elegansiforrnis Cookson, latest Maastrichtian

Proteaceae ?
Sapindaceae (Cupanieae
tribe)
Angiosperms of uncertain
or unknown affinities
B. verrucosus Cookson [Beauprea] latest Maastrichtian
Peninsulapollis gillii (Cookson) late Campanian-Danian-
Dettmann & Jarzen, P. askiniae Dettmann
& Jarzen, P. truswelliae Dettmann &
Jarzen, P. sp. [cf. Beauprea]
Cupanieidites orthoteichus Cookson & late Maastrichtian
Pike [Cupania]
Tricolpites spp.
Tricolpites striatus Couper late Maastrichtian
Tubulifloridites lilliei (Couper) late Campanian-M aastrichtian
Farabee & Canright 2
Forcipites sabulosus (Dettmann & late Campanian-early Danian
Playford) Dettmann & Jarzen
Forcipites sp. cf. F. longus (Stover & early Maastrichtian
Evans) Dettmann & Jarzen
Tricolporites pachyexinus (Couper) late Campanian-early Danian
Partridge
Tricolporites spp.
Rhoipites sp. cf. R. microreticulatus latest Maastrichtian-Danian
(Harris)
Rhoipites spp. Maastrichtian-Danian-
Polycolpites langstonii Stover late Campanian-Danian-
Tetracolporites verrucosus Stover late Campanian-Maastrichtian
Garnbierina rudata Stover late Campanian-late Maastrichtian
Stellidiopollis annulatus Dettmann late Campanian-Danian
& Hedlund; Stellidiopollis sp.
Triporopollenites sectilis Stover. late Campanian-late Maastrichtian
Triporopollenites spp.
Echitriporites spp. late Campanian-Danian
" Echitriporites" sp. late Maastrichtian-early Danian
Quadraplanus brossus Stover early to late Maastrichtian
Amosopollis cruciform& Cookson & Balme early to late Maastrichtian

"Danian-" denotes pollen species range into younger sediments on Seymour Island.
1A leaf impression and dispersed cuticle of Lauraceae (G. Upchurch, pers. commun., 1988) also occur in the Cross Valley Formation.
2Dettmann and Jarzen (1988a)

( H u o n pine) restricted to western T a s m a n i a where similar to Microcachryidites antarcticus and Podo-

it grows in a cool temperate maritime climate with
rainfall greater than 1200 m m / y e a r ( C o l h o u n et al.,
1988).
Extant N o t h o f a g u s f u s c a g r o u p species and
m a n y other p o d o c a r p a c e o u s conifers are charac-
teristic o f southern cool temperate floras. N. menzi-
esii g r o u p and the conifers Microcachrys and
Microstrobos occur in colder, higher altitude
conditions. The latter two are small shrubs in
subalpine, notably wet areas o f N e w South Wales
and Tasmania. T h e y p r o d u c e trisaccate pollen
sporites spp. pollen (e.g. C o o k s o n , 1947; Milden-
hall, 1978) which occur rarely but consistently
t h r o u g h o u t the S e y m o u r Island section.
A high h u m i d i t y / m o d e r a t e to high rainfall
regime is accepted for the C a m p a n i a n - P a l e o c e n e
n o r t h e r n Antarctic Peninsula. This is based on
paleobotanical (Francis, 1986) and palynological
evidence ( D e t t m a n n and T h o m s o n , 1987; Askin,
1988a, 1989) and on climate models (e.g. Parrish
et al., 1982).
The late Maastrichtian was relatively w a r m in
110
the northern Antarctic Peninsula area (60 ° to 70°S
paleolatitude). There the climate at the end of the
Cretaceous was humid, mild and equable (Askin,
1989). This premise is based on the Seymour Island
occurrence of fossil pollen from plants whose
presumed modern analogues (e.g. Beauprea, B o m -
bax, Anacolosa, Cupania) grow in the tropics and
subtropics, mostly in humid equable conditions.
Many of these plant species grow today at middle
to high altitudes. Pocknall and Crosbie (1988) and
Dettmann and Jarzen (1988a,b) have documented
fossil pollen similar to pollen of living Beauprea
and other New Caledonia plants, emphasizing the
similarity of late Cretaceous-Paleogene mid to
high latitude vegetation to that of present day New
Caledonia. Extant N o t h o f a g u s brassii group species
also grow in New Caledonia, at 150-1350m
altitude, and form a distinctive cloud forest zone in
New Guinea, between about 750 and 3000 m (van
Balgooy, 1971). Pollen of this group occur
throughout the Campanian to Eocene of Seymour
Island.
Pollen of some of the "warmth-loving" (frost
sensitive) forms are extremely rare fossils on
Seymour Island (Askin, 1989)• Consequently, it is
difficult to establish their complete stratigraphic
ranges with certainty. Perhaps significantly, they
occur in late Maastrichtian sediments and have not
been found in the Paleocene on Seymour Island.
Two of these species, Beaupreaidites elegansiformis
and B. verrucosus, reappear in Eocene sediments
on Seymour Island.
The Seymour Island terrestrial palynoflora
reflects changes in accessory angiosperm compo-
nents, but changed little in its dominant compo-
nents from the Campanian into the Paleocene.
P. m a w s o n i i continued as the dominant form and
proportions of other conifer and N o t h o f a g u s pollen
and common cryptogam spores, although showing
some fluctuations, did not change significantly.
Climatic changes (circulation, precipitation and
temperature changes) during this interval probably
resulted in altitudinal shifts of vegetation zones.
This could have produced grossly similar palyno-
floras, although elements at the temperature and
moisture extremes may have been lost to the
vegetation and consequently to the palynomorph
record.
R.A. ASKIN
Cretaceous/Tertiary boundary
The pattern of last and first appearances for
angiosperm pollen species across the Cretace-
ous/Tertiary (K/T) boundary on Seymour Island is
m
m •
.0
+1oo- 0
O0
+50'
LL
0 T +
--0-
0 35
t.-I species
rn
-50- (1)
"1o
N
Q.
-100-
Fig.2. Pattern of last and first appearances of-angiosperm
species over the Cretaceous/Tertiaryboundary. Sixteen addi-
tional specieshave already disappeared below this intervaland
35 speciescontinue through. Pollen speciesare: 1. Tricolporites
sp.2, 2. Bombacacidites sp., 3. Cranwellia spp. 4. Echitriporites
sp.2, 5. Gambierina rudata, 6. Cupanieidites orthoteichus, 7.
Tubulifloridites lilliei, 8. Triporopollenites sectilis, 9. Proteaci-
dites sp.4, 10. Tricolpites sp.6, 11. Cranwellipollis subpalisadus,
12. Triporopollenites sp.14, 13. Tricolporites pachyexinus, 14.
Nothofagidites sp.7, 15. Liliacidites sp.2, 16. Propylipollis sp.2,
17. Triorites sp.1, 18. Nothofagidites sp.14, 19. Liliacidites
kaitangataensis, 20. Stellidiopollis sp., 21. Bombacacidites
bombaxoides, 22• Tricolpites striatus, 23. Anacolosidites sectus,
24. Beaupreaidites elegansiformis, 25. "Echitriporites" sp. 1, 26.
Propylipollis retieuloscabratus, 27. Tricolpites sp.2, 28. Triporo-
pollenites sp.18, 29. Rhoipites sp.cf.R.microreticulatus, 30.
Tricolporites sp.4, 31. Beaupreaidites verrucosus, 32. Tricolpor-
ires sp.5, 33. Proteacidites sp.6, 34. Triporopollenites sp. 17, 35.
Proteacidites sp.8, 36. Triporopollenites sp. 1, 37. Triporopollen-
ires sp.19, 38. Nothofagidites sp.13, 39. Rhoipites sp.3, 40.
Tricolporites sp.3, 41. Rhoipites sp. 1.
SPORE AND POLLEN ASSEMBLAGES OF SEYMOUR ISLAND
shown in Fig.2. The interval shown spans 100 m
below and above the putative K/T boundary
located by dinoflagellate cysts (Askin, 1988b) and
other fossils (e.g. Macellari, 1985, 1988; Harwood,
1988; Huber, 1988). The cryptogam spore record
also shows some change. Three cryptogam species
(Polypodiisporites speciosus, Dictyophyllidites con-
cavus and Clavifera triplex) have their first appear-
ances within the 35m beneath the boundary
(Askin, in press), while other species appear above
the boundary. The conifer record shows no first or
last appearances in this 200 m interval. Many of
the angiosperm species that disappear near the
K/T boundary in southeastern Australia (e.g.
Helby et al., 1987) and New Zealand (e.g. Raine,
1984), including Tubulifloridites lilliei, Triporopollen-
ites sectilis and Quadraplanus brossus, also dis-
appear near the end of the Cretaceous on Seymour
Island, suggesting that a regional event/trend
affected the floras of the Weddellian Province.
In the late Maastrichtian the northern Antarctic
Peninsula climate was apparently humid, mild and
equable, but seems to have changed by the Danian,
based on the angiosperm pollen record. There was
a turnover in the angiosperm component in the 100
meters below and above the boundary (Fig.2).
Changes near the boundary may be related to
global environmental upheaval at the end of the
Cretaceous, but this is difficult to determine
because these transported, and probably reworked,
terrestrial palynomorphs might not be expected to
record a sudden event, especially if it was short-
lived.
A gradual trend of angiosperm disappearances
from the palynoflora is apparent from Fig.2,
although the ranges may have been extended
upwards by reworking. The gradual trend of first
appearances, especially noticeable immediately
below the KIT boundary, is more significant. It
suggests that some environmental change was in
progress, providing suitable conditions for immi-
gration or promoting speciation among angio-
sperms and cryptogams. Regression continuing
through the K/T transition may have been largely
responsible for changes in marine faunas and floras
(Macellari, 1988; Askin, 1988b). Regression may
also have directly affected land floras by providing
newly created lowland areas for new plant species.
111
A cooling trend may be partially responsible for
floral change observed near the boundary. This
concept is supported by the absence of so-called
"warmth-loving" angiosperm species in Danian
strata. No change in precipitation across the K/T
boundary can be inferred at present from the fossil
record.
Species diversity declines above the boundary,
although factors other than cooling may have
affected this parameter. In parts of the uppermost
L6pez de Bertodano (10) and basal Sobral (1-2)
Formations, restricted depositional environments
(e.g. estuarine) have resulted in palynomorph
assemblages with overwhelming relative abun-
dances of one or a few dinocyst species and
relatively few terrestrial palynomorphs. In the
overlying sandy sediments, palynomorphs are
often sparse, decreasing the likelihood of finding
many of the potential species.
There is some additional evidence of temper-
ature change. Oxygen isotope data from Seymour
Island benthic foraminifera suggest (Barrera et al.,
1987), however, that bottom shelf waters were
slightly cooler in the late Maastrichtian (between
4 ° and 8.5°C) than in the middle Maastrichtian
(5.5-9°C) and Danian (basal Sobral Formation,
5.5-10°C). The sampling intervals for analyzed
foraminifera and the broad estimated range of
temperatures may mask late Maastrichtian-basal
Danian temperature fluctuations. Oxygen isotope
data from O.D.P. Leg 113, Sites 689 and 690, on
the east side of the Weddell Sea (Stott and Kennett
et al., 1988), indicate relatively warm Late Cretace-
ous surface waters (16°C), then cooling immedi-
ately preceding the K/T boundary.
In conclusion, the late Cretaceous-early Terti-
ary vegetation of the northern Antarctic Peninsula
probably responded to climatic fluctuations with
altitudinal shifts of vegetation zones. This might
explain the continuity of dominant components in
the palynomorph record from the late Campanian
to Paleocene. Gradual disappearances and ad-
ditions of new taxa throughout this interval
probably resulted from the complex interaction of
climate changes, resultant altitudinal shifts of plant
associations and factors such as immigration,
speciation and transgression/regression.
112
Acknowledgements
V a l u a b l e c r i t i c i s m o f this m a n u s c r i p t b y t h e
r e v i e w e r s ( i n c l u d i n g D a l l a s C. M i l d e n h a l l ) a n d
S t e p h e n R . J a c o b s o n is m u c h a p p r e c i a t e d . I also
t h a n k M a r y E. D e t t m a n n and Elizabeth M.
Truswell for helpful discussion on particular
a s p e c t s o f t h e s e p a l y n o f l o r a s . S u p p o r t f o r this
p r o j e c t w a s p r o v i d e d by N a t i o n a l S c i e n c e F o u n d a -
t i o n g r a n t s D P P 8314186 a n d D P P 8716484.
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