The History of Neotropical Vegetation: New Developments and Status

Author(s): Robyn J. Burnham and Alan Graham

Reviewed work(s):
Source: Annals of the Missouri Botanical Garden, Vol. 86, No. 2 (Spring, 1999), pp. 546-589
Published by: Missouri Botanical Garden Press
Stable URL: http://www.jstor.org/stable/2666185 .
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THE HISTORY OF
NEOTROPICAL VEGETATION:
NEW DEVELOPMENTS AND
STATUS'

RobynJ. Burnham2and Alan Graham3

ABSTRACT
The isolationofSouthAmericafromCentralAmericaand AfricaduringtheTertiary
Periodlefta strongimprint
on
thefloraoftheNeotropics.SouthAmericanEocene through
Miocenefossilassemblages,bothpollenand macrofossils,
documenta rich tropicalfloraon the continentalmargins,and representsome of the onlydata on pre-landbridge
lowlandtaxa in SouthAmerica.LowlandMioceneflorasfromAmazoniaare remarkably
similarin theirhighdiversity
to Amazonianflorastodaybased on lists of dominantfamilies.Recentgeophysicaldata on the upliftof the northern
ofmontaneforestsin Colombia
Andesshowa strongcorrelation
betweenupliftand thedevelopment
and diversification
and Venezuela.The emergenceof a continuouslandbridgeat 3 Ma betweenCentraland South Americais well
documentedand is demonstrated
by the arrivalof temperateelementsin SouthAmericanhighlandsand concurrent
appearanceof SouthAmericantaxa in CentralAmerica.There is no evidencefordisplacementof lowlandtropical
plants in South Americaby northern
immigrants,
whichappears to stand in contrastto the publishedrecordfor
mammals.The mix of taxa in extantMexican tropicalflorasderivedfromtropicalSouthAmerica,tropicalCentral
America,and fromremnants
of northern
tropicalEocene florasis strongevidenceforthe impactthatthe landbridge
through
thePanamanianisthmushad on theneotropical
flora.The earlyappearanceoflow-elevation
savannasis inferred
froman increasein grasspollen in the middlePliocene of Panama; however,widespreadsavannasare notindicated
by pollendata fromthe CentralAmericanregion.Rather,beginningin thelatestMioceneEpoch and continuing
up to
theQuaternary,
a mixoftropicalrainforest
and mixedtropicalwoodlandsis suggestedforthelowlands,based on pollen
evidence.Accumulating
data on temperature
and Quaternary
Periodspointsto lowchangesduringthe late Tertiary
latitudetemperature
fluctuations
of up to 60C. Proposalsof accompanying
widespreadrainfallfluctuations
are more
sea level
equivocal.Rainfallprobablyvariedregionally,
resultingin a mosaicofhabitatscontrolledbyrivermigration,
local dryness,and local uplift.Zones postulatedas refugiaprovidetestablehypotheses
fluctuations,
usingneoecological
and paleoecologicaldata. The paleoecologicaldata to testthesehypotheses
and spatially.
are stilllimitedtaxonomically
It is important
to stressthattheeffectofthe isolationofSouthAmericanneotropicalflorashas notbeen erasedin the
3 millionyearssince theirconnectionwithCentralAmerica.New data frommiddleand late Mioceneflorasin South
Americawill be criticalin determining
the degree to whichthe compositionof South Americanflorashas been
influenced
ofplantsfromthebetter-known
by immigration
CentralAmericanarea to thenorth.

The neotropicsextendgeographically
fromthe fromthese extra-tropicalsources are present. There
Tropicof Cancer to the Tropic of Capricorn,in- are as many reasons for neotropic patterns of dicludingenvironments
as diverseas drydesertor versityand distributiontoday as there are different
humidrainforest.
Mean annualtemperatures
range landformsand climatic regimes. If these differences
fromover30'C to as low as 10'C. Elevationsfrom are played out over the course of the late Mesozoic
sea level up to well over6000 m are included.This and Cenozoic, when data fromfossil deposits can
wide rangeof climateand topography
has a pro- informus about the sequence of changes that has
foundeffecton the compositionand structureof taken place, the history of neotropical vegetation
neotropicalvegetation.
The area also has a distinc- takes on a dizzying complexity.
tive geographicoutline. It is roughlyhour-glass
We approach this review of the vegetational hisshaped,withthe narrowisthmusof Panama com- tory of the Neotropics by outlining four important
posingthefragile,primarily
lowlandconnection
be- events (Fig. 1). We initiallytreatthe two major land
tweenlargenorthern
and southernland masses.In areas separately, because for much of their history
addition,boththenorthern
and southern
neotropics they were indeed isolated fromone another. Then,
lie contiguousto subtropicaland temperateareas we summarize the interactions that postdate the
to the northand south:no barriersto immigration connection and treat selected aspects of the Neo1 We are grateful
forcarefulreviewof earlierversionsof this manuscript
by Paul A. Colinvaux,PeterR. Crane,
David L. Dilcher,and twoanonymous
reviewers.We also extendour appreciationto Bruce D. Pattersonand David S.
Webbfordiscussionof and comments
on patternsof mammaliandiversification
and migration.
2 Museumof Paleontology,
University
of Michigan,Ann Arbor,Michigan48109-1079, U.S.A.
3Department of BiologicalSciences,KentStateUniversity,
Kent,Ohio 44242, U.S.A.

ANN. MISSOURI BOT. GARD. 86:

546-589. 1999.

Burnham& Graham
Historyof NeotropicalVegetation

Volume 86, Number2

1999

A PARTIAL TIME SCALE

ERA

PERIOD!
SUBERA

EPOCH!
STAGE
____

O
N
O

(CRETACEOUS

___

___

w
w

Present

OLIGOCENE
EOCENE

a-

PALEOCENE

UPLIFT

56.5

CAMPANIAN

Cl)

LANDBRIDGE
EXISTS

34

MAESTRICHT.

C)

UATERNARY
CLIMATE

23.3

O
N

TO PRESENT)

Million NEOTROPICAL
Years
EET
Before
EVENTS

HOLOCENE
QUATERNARY
PLEISTOCENE 0.01
1.64
z
PLIOCENE
w
5.2
O
w
MIOCENE
>-Z

547

65

ISOLATION

74

SANTONIAN
CONIACIAN 88.5
TURONIAN
0

CENOMANIAN
ALBIAN

~~~~~97

APTIAN
to Presentshowingthe foureventsdiscussed in the text.Note that
Figure1. Time scale forthe mid-Cretaceous
the eventsare nottemporally
distinctfromone another.

tropicsas a whole up to the present.The unique lished fossil florasfromCosta Rica to Bolivia,
in Appendix1.
historyof the vegetationof the Neotropicsis, in whichare referenced
large part,due to the geographicisolationof its
southernhalf duringmostof the past 90 million A BRIEF HISTORY OF THE ACCUMULATION OF
years,subsequentupliftof the Andes, and inter- MACROFOSSIL AND MICROFOSSIL
actions betweenthe northernand southernele- DATA IN THE NEOTROPICS
ments.
Amongthe oldestreportsof plantfossilassemWe reviewdata on the pollen and macrofossil blagesfromtheNeotropicsare theworksofT. Wolf,
recordsfromthelate CretaceousPeriodup to about a geologistwhomappedareas ofsouthernEcuador
5000 years ago, when climatesand physiography in the late 1800s in companywithG. vom Rath
tookon essentiallymodernaspects.We addressthe (Wolf& vomRath,1876). His workon paleontology
Neotropicsas geographicallydefined:from230 and geologyin southernEcuador coincidedtemSouth to 230 Northlatitude.At the southernex- porallywiththeexplorations
on extantplantbiology
tremewe have includedfossilflorasfromBolivia, of Richard Spruce in the Amazon and Andes.
buthave excludedflorasfromArgentina,
Paraguay, Wolf'sreportsoffossilsfromthe inter-Andean
baand southernBrazil. At the northern
extreme,we sins of southernEcuador werefollowedby H. Enhave includedfossilflorasfromMexicoup to 230 gelhardt,who published several taxonomictreatNorthlatitude.Ourreviewofthehistory
oftheveg- mentsof plant fossils from1887 through1895.
etationin this broad geographicarea incorporates Fromabout 1915 through1945, E. W. Berrypuball the publisheddata known,insofaras it can be lished extensively
on neotropicalpaleobotanywith
integrated
intodatabaseson plantmacrofossils
and workson macrofossilflorasfromMexico through
microfossils.
Figure 2 showsthe locationof pub- Patagonia (Berry,1921a, 1929a, 1945a). Berry

548
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exnumberof knownmodemtaxa. His firsthand

periencein SouthAmericaappears to have been
ofEocene fosquite limited.Berry'sidentifications
sil plantsfromthe UnitedStates may be at least
60% in error(Dilcher,1974), and it is also a concern thatpollen samplesprocessedfromthe same
Costa Rican localities (Graham,1987) turnedup
none of the genera identifiedby Berry(1921a).
C/)12
was domthepaleobotanicalliterature
Nonetheless,
Z
POLLEN
0
inatedby Berry'spublicationson theNeotropicsup
0 MACROFOSSILS
to the timeof his death(Appendix1).
C-)
AfterBerrytherewas a relativelull in publicam
tion activity(Fig. 3) untilpalynologybecame an
establishedresearchtool in tropicalAmerica.In
0 4
worksof T. van
1954 the firstof the monumental
uJ
L2L
m2
using
der Hammenon vegetationreconstruction,
oftheColombianAndes,was published
palynology
D
O C O C\j N- C\j N- C\j N- C\j N- C\j N- C\j N- C\j N- C\j NCO C- 0(van der Hammen,1954). Some of thisworkused
makingcomparitaxonomy,
form-generic
artificial,
YEAR
difficult.
However,otherresons withmacrofossils
Figure3. Publicationactivityin neoptropicalpaleo- search was focusedon creatinga pollen zonation
botanysince 1876. Publicationsare coded withrespectto forthe High Plain of Bogota.Fromthe dominant
the dominantorgan reported:all macrofossils(leaves,
of vegetationzones were
fruits,seeds, wood) are grouped.Publicationslisted are taxa, reconstructions
reportsof localitiesand fossilplantremains;no achieved and directlyrelatedto the upliftof the
primary
Andes.
northern
summaries
or reviewsare included.
publishedmorethan55 paperson SouthAmerican
paleobotanyalone, detailingthe systematicaffinialvegetation,
ties of the florasand reconstructing
titude,and paleoclimate.AlthoughBerry'streatmentsof the floraswere classic forthe time,his
small
werebased on a comparatively
identifications

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Volume 86, Number2

1999

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Burnham& Graham
Historyof NeotropicalVegetation

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The potentialfor assessing the natureof past DISTRIBUTION OF THE DATA IN TIME AND SPACE
ofmacdependsuponthemethodologies
vegetation
In Figure2 we showthe geographicaldistribuEach has
rofossilpaleobotanyand paleopalynology.
of data used in this review.Localities from
tion
an
Pollen
provides
and weaknesses.
its strengths
fossilflorashave been described,eitherwith
which
of manycanopytrees,shrubs,and uninventory
cenor floristic
ofvegetationreconstruction
goal
a
herbaceousplants,identifiablemostlyto
derstory
time
the
for
coded
geological
and
were
plotted
sus,
larger-scale
genus,and is usefulforreconstructing
are
these
on
shown
maps
Sites
represented.
period
of
kilometers
of
square
(tens
patterns
vegetational
Apin
position
stratigraphic
and
country
by
listed
to a singlesample).Macmaycontribute
vegetation
in the
ofsitesmentioned
rofossilsprovidean inventorymostlyof canopy pendix1. The distribution
Antilles
the
and
America,
Central
in
Mexico,
text
allow
to species,and
treesand shrubs,identifiable
of vegetationin the imme- are presentedin publicationsby Graham(1988c).
detailedreconstruction
diate vicinityofthe depositionalbasin; less thana In SouthAmericathereis a clear preponderance
square kilometerof vegetationusuallycontributes of localities on continentalmargins.Amazoniais
sites,and it is
poorin pre-Quaternary
to a single fossil assemblage (Burnham,1994). particularly
Clearly,the integrateduse of both methodologies importantto continueexplorationfor additional
createsthe highestresolutionpictureofvegetation palynologicalor macrofossilassemblages in the
and vegetationalchange.Althoughpairingofsam- rich Cenozoic deposits (Tschopp, 1953; Duarte,
ples fromlocalitiesis a logicalgoal,itis onlyrarely 1972, 1983, 1985).
We have also displayedthe data for northern
thatsuch studieshave been carriedout in paleoAmerica in two categories:pre-1965 and
South
botany(Hollick,1928; Graham& Jarzen,1969; Le(Figs. 4-8). This distinctionis entirely
post-1965
1986;
Dilcher,
&
Farley
1972;
opold& MacGinitie,
chosen only on the basis of a visible
Farley& Wing,1989; Taggart& Cross,1990; Skog arbitrary,
& Dilcher,1994; Willardet al., 1995; Wijninga, break in publicationactivityat 1965. In addition,
datingwas beingwidelyappliedbythis
radiometric
1996a).

550

time,and some tropicalCenozoic florascould be

dated independently
of the estimatesbased on the
fossilassemblages.Thus thetwocategoriesalso reflectgreaterpossibilityforindependentage confirmationin the morerecentlypublishedworks.

Annals of the
MissouriBotanical Garden

of continental drift (Sullivan, 1974; Graham,

1988c), a situationthatdid not encouragecritical
comparisonofthefossilflorasofthetwocontinents.
of
In fact,theorieson the originand development
the biogeographic
affinities
of the SouthAmerican
florawerederivedmorethroughstudyofthe modem flora(Raven & Axelrod,1974; Gentry,
1982a,
FOUR MAJOR EVENTS STRUCTURING
b, 1990; Simpson & Neff, 1985) rather than
NEOTROPICAL VEGETATION
ofpaleobotanists
theefforts
ofthetime(but
through
EVENT I: ISOLATION
see reviewsbyTaylor,1991, 1995). Thus,themeager historyof comparisonof floristicpatternsbeThephysicalsetting
tweenCentraland SouthAmericamay be due to
oftheflorasand an earlylack of
Geophysicaland biologicalevidencenow indi- thehighdiversity
cate thatrifting
betweenAfricaand SouthAmerica acceptanceofthe conceptsofcontinental
drift.
was underway by about 95 mya (Pitmanet al.,
1993; 106-84 mya,Goldblatt,1993). By85 million Floristic similaritybetweennorthernand
yearsago, it is estimatedthata seawaycoveringas southernNeotropics
muchas 100 longitude(up to 800 km) existedbeFossil angiosperm
florasin northern
SouthAmertweenthe two southerncontinents.
Plant commuica older than65 mya are few(Fig. 4). More are
nities in South Americaof this age are too fragavailable forassessingthe degreeofisolationfrom
mentaryto providemuch evidence on the degree
fossilflorasof earlyTertiary
age (Fig. 5). The maofisolationoftheflorasofthetwocontinents.
Howjorityofthereported
florasare fromColombia,Venever,fromotherregionswe knowthatangiosperms
ezuela, and Ecuador,withoverhalfpublishedbewerediversified
by thistime(Muller,1981, 1984;
fore 1965. To the north,well-preservedearly
Winget al., 1993), and severalmodemfamiliesare
Tertiarypollen florasare reportedfromPanama
recognizable(Wolfe& Upchurch,1987; Friis &
(Graham,1985, 1999a) and Mexico(Martinez-HerCrepet,1987). The ecologicaldominanceofangionandez et al., 1980). The percentsimilaritybespermsin mosthabitatsbecomesapparentafterthe
in Early to Middle Eocene
tween palynomorphs
Cenomanian(Wing& Tiffney,
1987a, b; Wing et
pollen floras fromPanama and northernSouth
al., 1993).
Americawas calculatedby Graham(1992) and is
At the same timethatAfricaand SouthAmerica
reportedherein Table 1. The lowdegreeoffloristic
were separated,South Americawas also isolated
similarity
(2.6%) is in accordwiththephysicalrefromNorthAmericaby a marineportalthrough
constructions of the Panamanian landbridge
CentralAmerica.This conditionhad existedsince
(Coates et al., 1992; Coates & Obando, 1996), as
the inceptionoftheTethysseaway(EarlyJurassic)
ofmamwellas withthedata on thedistinctiveness
and continueduntil the emergenceof the Panamalian faunas at thattime.The taxonomicdetermanianLandbridgein theLate Pliocene.Thus,notminations
made on theleafflorasare eitherlimited
the possibilityof small-scalechance
withstanding
in numberor too unreliableto reconstruct
vegetadispersalevents,SouthAmericafromabout 100 to
Eotiontypesin thePaleocene. Recently,
however,
about3 millionyearsago was an island continent
cene mangrovevegetationflankedby inlandpalm
(McKenna,1980; Simpson,1980; Marshallet al.,
1982; Simpson& Neff,1985; Webb, 1985; Marshall, 1988). The isolationof South Americaatof palynomorphs
betweenCentral
Table 1. Similarity
tractedthe attentionof vertebratepaleontologists and SouthAmericafromtheEarlyEocene Epoch through
because fossil mammalsin South America,older Quaternary
Period.
thantheconnectionwithNorthAmerica,wereeasfromtheirnorthern
ily distinguished
counterparts.
of
Similarity
between
palynomorphs
Paleobotanists
wereslowerto recognizethedisconCentralAmericaand
tinuities,perhapsbecause the extantangiosperm
Time period
SouthAmerica
floraofnorthern
SouthAmericaand CentralAmerEarlyto MiddleEocene
2.6%
ica was bewilderingly
Hemidiverseto Northern
10.7%
EarlyMiocene
sphere plant biologists,but also because many
MiddlePliocene
8.9%
plantcollectionsfromSouthAmericawerelimited
-Landbridge
establishedand thisprovidedfewopportunities
forcomparison.
29.7%
Quaternary
In addition,Berrywas notan adherentofthetheory

Volume 86, Number2

1999

Burnham& Graham
Historyof NeotropicalVegetation

551

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forestshas been recon- geographicbarrieris responsibleformanyof the

and dicotyledon-dominated
coastof observedpatterns.
structedbyRull (1998a) fromthenorthern
SouthAmerica.
familydiversity
Fossil and modern
distributions
modern
in
recognized
Disjunctions
Duringthe period of isolation,northernSouth
harboreda richassemblageofangiosperm
America
Terthe
during
America
South
of
The isolation
tiaryPeriod can also be inferredfromthe distri- families.The high diversityof Amazonianforests
(1992), todayis due in partto recentspeciationand imbutionofextanttaxaas presentedbyGentry
of taxa fromCentralAmericastarting3
Wendt(1993), and Hammel and Zamora (1990, migration
fora groupofplants millionyears ago, but the high diversitywithin
1993). The predicteddiversity
neotropicalfamiliesmostlikelyis due
of prominent
isolatedin South Americauntil establishment
historyofmanytaxonomic
thelandbridgewouldbe a highnumberof species to the longevolutionary
fossil
AlthoughTertiary
America.
in
South
groups
wideperhaps
few,
a
only
in SouthAmerica,with
spread,taxa extendingintoCentralAmerica.This florasfromAmazoniaare few,none of themcomformanygenerabyGentry pletelydocumented,and mostfrommoderatepawas documented
pattern
(1982b) and morerecentlyforSerjania sect. Pla- leoaltitudes(probablyclose to 1000 m), fossilas(1993). Such a pat- semblages of Miocene deposits fromsouthern
byAcevedo-Rodriguez
tycoccus
tern,however,mightalso derivefromnarroweco- Ecuador (Berry,1929a, 1945a) do reveal familyFossil taxawereassessed onlyat the
taxa and wider level diversity.
logicalamplitudesin the restricted
by Berry
the identifications
because
level
family
widein
the
geographically
ecologicalamplitudes
taxa.Nonetheless,in manyofthecases cit- have been repeatedlyquestionedoverthe past 40
ranging
under
ed by Gentry(1982a), the rangesof SouthAmeri- years.Florasfromthesebasins are currently
Beron
research
our
and
of
us
one
(RJB),
by
study
across
can taxaextendnorthintoColombia,butnot
shows
taxonomicdeterminations
theIsthmusofPanama. This suggeststhata recent ry's family-level

552
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of isolation
thattheyare reasonablein about70% ofthecases. Endemismas an effect
Four modernlowlandtropicalfloraswerereported
A patternexpectedfromthe isolationofa land1990) in whichthe percentageof species
(Gentry,
mass fora considerableamountof timeis a high
familiesare listed,and thesecan
in 20 angiosperm
level of endemism.This is well documentedon isof species in the
be comparedto the proportion
of Madalands. For example,the island continent
same familiesoffossilplantsfromEcuador(Fig.9).
gascarincludesclose to 10,000 endemicplantspethemostcommon
Atthiscrudelevel ofcomparison,
cies due in partto its isolationfromAfricasince
familiesin thefourmodernAmazonian
angiosperm
(Takhtajan,1986). The 3 milthe mid-Cretaceous
forestsare also representedin the fossil assemlion yearsofcontactbetweenCentralAmericaand
blage. Similarly,Rull (1998a) indicatedthat the
northernSouth America has certainlydecreased
Veneof northern
Eocene mangrovecommunities
theextentofendemismin bothareas,buthighento
and composition
zuela were similarin diversity
demismin some groupsstill can be documented.
other equivalent-agemangroveflorasworldwide.
Table 2 listsgenerawithat least threespecies that
Hoorn(1994a) reportedMiocenepalynologicaldiprobablyevolvedin SouthAmericaand are endemversityin AmazonianColombiathatis at least as
fosic theretoday.GeneraknownfromtheTertiary
high,if not higher,than moderndiversityin the
sil recordin South Americaare indicated.Like&
area today(see also discussionby Hooghiemstra
wise, Wendt(1993) listed genera that may have
van der Hammen,1998). Lundbergand Chernoff
in CentralAmericaor Mexico and are
diversified
(1992) and Lundberg(1997) reportedthatfreshendemictheretoday.
in Middle Miocenedepositsof
waterfishdiversity
Colombiawas probablyas richas in thefishfaunas
EVENT II: UPLIFT AND PHYSIOGRAPHIC CHANGE
fromAmazonianbasinstoday.Mammaliandiversity
The second importanttime in northernSouth
was also high beforethe interchange(Marshall,
Americais the periodfromabout 15 millionyears
1985; Marshall& Cifelli,1990).

Volume 86, Number2

1999

Burnham& Graham

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ago to thepresent.Notethatthisphase, including

upliftoftheAndesand changingdrainagepatterns,
periodoflandbridgeconoverlapswiththeyounger
nectionand also withthe olderperiodofisolation.
in interpreting
thepresentvegPartofthedifficulty
etationin SouthAmericaderivesfromeventsthat
overlap.This underscoresthe care that mustbe
patternsnotedin moderndistakenin attributing
in climate.
tribution
to recentfluctuations
Altitudinal
changes
The major period of upliftin northernSouth
Americaoccurredin the MioceneEpoch, although
thereis strongevidenceforupliftin thecentraland
southernpart of South Americaearlier than the
MioceneEpoch (Jordan& Alonso,1987; Sempere
to deteret al., 1990). Exact altitudesare difficult
pairedlowminebecause thisrequireslatitudinally
land and uplandequivalent-ageflorasthatcan provide lapse rates or enthalpyestimates(and thus
altitudes:Wolfe,1992; Forestet al., 1995; GregoryWodzicki,1997; Gregory& McIntosh,1996). Despitethe large numberof florasstudiedby Berry,

...............

dates,and manyappear
feware tiedto radiometric
to be fromareas of moderatepaleoelevation.Lowland coastal or lowlandAmazonianflorasexistin
Colombia,Ecuador, Peru, and Brazil, but these
have notbeen studiedor are nottiedto well-estabsections.
lished biostratigraphic
in western Colombia
The paleophysiography
as an elongatedpeninsula
has been reconstructed
witha mountainousbackbone. Elevationsduring
Andes
the middleMiocene Epoch in the northern
have been estimatedbypalynologicalrecordsfrom
the High Plain of Bogota'as being less than 700
m, while by the late Miocene Epoch the area is
estimatedto have been close to 1000 m (Wijninga,
1996b). Middle Miocene sedimentsin centralColombia preservethe exquisite La Venta mammal
fauna, interpretedto indicate lowland tropical
conditions(Kay & Madden,1997) probablybelow
500 m. Situatedon the easterncordilleraof the
ColombianAndes, the mammalfossils and their
indicatethatupliftbesedimentary
environments
gan as early as 12.9 mya (Guerrero,1997). The
La Ventasites would be ideal forcalibratingup-

554

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of Quater

Distribution

fossil plant localiti

of published

C/)

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W

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41"

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Volume 86, Number2

1999

land florasif fossilleaf florascould be recovered

fromthis volcanicallyinfluencedterrane.Fossil
wood fromthe area was reportedby Schbnfeld
(1947). Fish faunas datingfromthe middle Miocene Epoch and preservedin the presentMagdalena Valleyof Colombiadocumenta strongafto currentAmazonianfreshwater
finity
fishfaunas.
This indicatesthatthe Eastern Cordilleraof Colombia had not created sufficient
barriersto migrationoffishuntilat least 11 mya (Lundberg&
Chernoff,
1992).

Burnham& Graham
Historyof NeotropicalVegetation

555

Hammen,1998). The palynologicalworkof Hoorn

(1994a, b) and Wijninga(1996a, b) addressedthe
easternand centralregionsof Colombia,respectively.Hoorndocumenteda varietyof habitatsinand dicludingpalm swamps,riparianvegetation,
versetropicalforestsin easternColombia.Flooding
was common,and the majorityof sedimentcame
fromthe east. She also presentedstrongevidence
forvegetationchange due to upliftin the Andes
duringthe middleto late Miocene(Hoorn,1994b).
Wijninga,whoseworkinvolvedsedimentsofmiddle
Miocene throughlate Pliocene age, indicatedthat
Drainage and physiographic
change
in the presentHigh Plain of Bogota'lowlandconditionsexistedthrough
theend oftheMioceneEpofthe physiography
Reconstructions
ofnorthern
och. Lowland forestsdeveloped on well-drained
South America presentedby Hoorn (1994a, b,
floodplains.
Higherelevationssupportedonlylower
1995) and by Vonhofet al. (1998) and synthesized
montaneforests.By theearlyPliocenesub-Andean
by Kay and Madden(1997) showthatin the early
taxa appearedon thepresentHighPlain ofBogota',
Miocene Epoch, the WesternCordilleraof Colomalong withthe pollen of a fewhigh-Andeantaxa,
bia was highenoughto deflectsedimentin an eastwhich were interpretedas comingfromnearby
ward directionand potentiallyisolate the current
highlands.Wijninga(1996a) documentedthe proChoco region.The major sedimentsource to the
gressiveupliftof the Andes throughthe Pliocene
Amazon Basin was fromthe moderatelyuplifted
usinga combinedpollenand macrofossil
approach.
Guyana Shield (Fig. 10). A controversial
portal
Analysisof the upper stratigraphic
sequence was
(Maraflon
Portal)in thearea ofGuayaquilhas been
presentedby Hooghiemstra
(1994), whodocumentproposed(Nuttall,1990; Hoorn,1994a), butbythe
ed the arrivalof northern
temperatetaxa (see bemiddleMioceneEpoch, theAndes in southern
Eclow).
uador probablyno longerallowedwestwardtransThe effectsofthe changingdrainagepatternsin
portof sedimentsfromthe Amazon Basin to the
northern
SouthAmericaare also preservedin strata
Guayaquilarea (Vonhofet al., 1998). Instead,priat the mouthsof the majorrivers.One such study
withsedimarysedimentfluxwas northeastward,
(Diaz de Gamero,1995) indicatedthatin theearly
mentsdirectedtowardthemajordepositionalbasin
River
throughmiddle Miocene the proto-Orinoco
offthe coast of presenteasternVenezuela. Curbuilta deltain northern
Venezuela,east oftheMathe Andes providethe bulk of sedimentsto
rently,
racaibo Basin. Throughupliftof the EasternCorthe Amazonand Orinocosystems,withthe largest
dillera of Colombia,the riverwas deflectedprosedimentflux now directedeast to the Amazon
gressivelyeastwardduringthelate middleand late
mouth,ratherthannorthtowardthe Orinoco.
Miocene.This upliftto thewestofthedepositional
The simplified
physiography
presentedheresugcenterisolatedthe florasof westernColombia,as
geststhatthe middleMiocene is the timeduring
well as fragmented
and diversifiedhabitatsthat
whichthedistinctive
natureoftheflorain theChopreviouslyexisted as contiguoustropicallowland
co regionbegan its development.
Habitatdiversity
swamps.
in the Choco regionwas probablyhigherin the
mid-Miocenethan suggestedby Gentry(1982a),
whichwouldgive a muchgreaterage to the ende- Evidence fromplant macrofossils
mismdocumentedtheretoday.The Choco,includThe record of Miocene plant macrofossilsin
ing coastalEcuador,was isolatedfromtheAmazon
northern
SouthAmericais seeminglyrich,with50
by the risingAndes and fromPanama by thedeep
publishedreports.However,muchofthisworkwas
trenchoffthe northwestern
coast of Colombia.
completedpriorto 1960, and lacks a modernsystematicand plate tectonicperspective.One of us
Evidencefrompalynology
(RJB)has recentlyinitiatedmacrofossil
plantstudCertainlythe most useful paleobotanicalre- ies in Ecuador and Bolivia (Burnham,1995a, b).
Ecuadorand two
search in northern
SouthAmericain the past 50 ThreeMiocenebasinsin southern
The threeEcuyearsis thatofT. van der Hammen,H. Hooghiem- in Bolivia are underinvestigation.
ofAm- adorianbasinseach include55-75 species offossil
stra,and theirassociatesat the University
sterdam(summarizedin Hooghiemstra
& van der plants, of which most are dicotyledonousangio-

Annals of the
MissouriBotanical Garden

556

genera, still
Table 2. Selected Gondwanan-derived
primarilyendemic to South America today.Estimated
numberof species per genusis indicatedin parentheses.
Onlygenerawithat least threespecies are listed.* indicates Tertiary
fossilrecordin SouthAmerica.
Trees
Amaioua(25)
*Apeiba(10)
Campomanesia(80)
Catoblastus(17)
(30)
*Courssapoa
(17)
Crematosperma
(5)
Cyclolobium
Ecclinusa(21)
(3)
Geoffroea
Herrania(20)
Hexachlamys(3)
*Humiria(4)
Jessenia(6)
Leonia (6)
(5)
*Loxopterygium
Parapiptadenia(3)
Peritassa(14)
Ptilochaeta(5)
Seguieria(6)
Schinopsis(7)
Siparuna(150)
Socratea(5)
*Thinouia(12)
*Ticorea(3)

Lianas and vines

Clytostoma
(9)
Mansoa (15)
Maripa (19)
Selysia(3)
Siolmatra(3)
Apondandra(4)
Dicella (6)
*Trigonia(24)

EARLYMIOCENE
Shield
Guyana

BrasilanSil

MIDDLETO LATEMIOCENE

GuyaaSil

ShIeld
Brasilian

sperms. Species richness is high in some localities

(up to 33 species). Over 80% of the species in each
basin bear entire-marginedleaves, and the leaf size
a
~~PRESENT
is about 35% notophyll,40% microphyll,and the
remainder evenly divided between very small and
Shield
Guyana
very large leaves. Provisionally identified angiosperms fromthe Cuenca Basin are: Loxopterygium
(Anacardiaceae), Tipuana (Fabaceae: Burnham,
1995a), Coussapoa (Cecropiaceae), Lauraceae, Malpighiaceae, and Arecaceae, while the Nabon Basin
includes Tipuana (Fabaceae), Roupala (ProteaShield
Brasilian
ceae), Sapindaceae, Clusiaceae, and Melastomataceae. The Loja Basin was studied by Berry(1929a,
1945a); taxa fromthatbasin verifiedby recent work
include Coussapoa (Cecropiaceae), Trema (Sterculiaceae), Tipuana (Fabaceae), Ruprechtia (Polygonaceae), Meliaceae, Lauraceae, and Malpighiaceae.
Directionof major
Directionof minor
Marine incursion
The Miocene basins of southernEcuador are well
sediment influx
sediment influx
suited to provide informationon the rate of uplift
and degree of influence of a rain shadow in the
sources,highlandareFigure10. Major sedimentary
northern Andes during the middle Miocene. All as, and seawaysin Early Miocene,Middle to Late Miothree floras have been tied to radiometric dates cene, and PresentSouthAmerica.FromHoorn(1994b).
rangingfrom13 to 10 mya (R. H. Madden, unpublished data). Mean annual paleotemperature indi-

Volume 86, Number2

1999

Burnham& Graham
Historyof NeotropicalVegetation

557

offlora andfauna
cated by leaf marginanalysisvariesamongtheba- Interchange
deposits
sins from22 to 280C. All plant-bearing
Of particularbiogeographicimportanceforthe
of small leafletsof Fainclude a large proportion
floral
and faunalinterchangeis the timingof the
baceae (up to 30% of the species), whichmayinformation
ofthePanamalandbridge.This has been
dicate a seasonallydryclimate.Of the threesites,
sources:
fromthreeindependent
recently
estimated
Loja has the largest-leavedspecies indicatingrelmaof easternPacific/Caribbean
of provincialization
ativelywetterconditionsbased on correlations
et
al.,
Coates
1996;
&
Obando,
(Coates
biotas
rine
in modemfloras(Wilfet
leaf size and precipitation
ofterresJacksonet al., 1996), interchange
1992;
al., 1998). Like today,Loja mayhave been to the
trialmammalsbetweenNorthand SouthAmerica
east oftheAndes,whereastheothertwobasinsmay
(Stehli& Webb,1985; Webb& Rancy,1996), and
havebeen betweenrisingparallelchainsoftheAnon eitherside
betweenpalynomorphs
thesimilarity
des. This easternpositionwould have diminished
timesduringthe Cenoof the Isthmusat different
the effectof a rain shadowon thefloraofthe Loja
zoic (Graham,1992).
basin.
Throughthe middleMiocene (16-15 Ma) there
and its effecton vegChangesin physiography
were marinepassages across present-dayCentral
etationare well documentedin sectionsfromCoAmerica,and deep-watersedimentsand biotasare
lombia,butnewevidencefromthePebas formation
especially evidentbetweenthe southern(eastern)
in easternPeru (Rdstinenet al., 1987, 1995) indiAmericaand the cordilterminusof proto-Central
cates thatchanges datingto the middle Miocene lera of northwestern
SouthAmerica(Fig. 11). By
northern
SouthAmerica.In- the late Miocene (7-6 Ma) marineportalswere
are presentthroughout
deed, as notedby Guerrero(1997), thetimeofup- morerestricted
(Fig. 12), and by the late Pliocene
liftof the Eastern Cordilleraof Colombiawas a (-3 Ma) theyexistedonlyacross the easternand
which included the westernextremesof the Isthmus(Fig. 13). Sediperiod of global importance,
sea- mentological
closureof the east end of the Mediterranean
provincialism
and marineinvertebrate
way; the collisionof the Australianplate withIn- date the establishment
of an essentiallycomplete
donesia,thusclosingthefaunalgap recognizedas structuralconnectionbetween Northand South
hiatus Americaat between3.5 and 3.1 Ma (Coates et al.,
Wallace's line; the deep sea sedimentation
at 12.0-11.8 Ma reportedby Keller and Barron 1992; Coates & Obando, 1996). Sometimeafter
(1987); and a majorcoolingeventthatexpanded 3.5-3.1 Ma theinterchange
ofterrestrial
organisms
the AntarcticIce Cap (Woodruff
et al., 1981). In- lackingmeans forlong-distancedispersalwas facreases in the seasonalityof precipitationhave cilitatedby the development
that
of land corridors
been citedfrommanyareas,whichwouldhavesub- were elevated above the low-lyingcoasts and
at thistime.
increasedhabitatdiversity
stantially
waifdispersaleventspriorto
Although
marshlands.
3.5 mya have been noted(Reig, 1980; Gingerich,
1985; Gentry,1992; Marshall& Sempere,1993;
EVENT III: CLOSURE OF THE ISTHMUS OF PANAMA
Webb & Rancy,1996), thereis no conclusiveeviPhysicalchanges
corridor
existed
dence thata continuousterrestrial
betweenCentralAmericato SouthAmericapriorto
betweenNorth the late Pliocene,about3 to 3.5 mya.
ofland connections
The formation
and SouthAmericahad profoundclimatic(Burton
Recentsummariesof the mammalianfossilrecet al., 1997) and biogeographicconsequencesfor ord provideimportant
information
on the climates,
Priorto the kindsof habitats,and the timeof appearanceand
thefloraand faunaofbothcontinents.
middleMioceneat -15 Ma tropicalmarinewaters expansionof the terrestrial
corridors.
First,the afinterchangedbetween the Atlantic and Pacific finitiesof the Earlyand MiddleTertiary
faunasof
Oceans, and there was only weak marine-water CentralAmericaare almostexclusivelywithNorth
northward
via the GulfStream.At -15 America,whiletherelationship
transport
ofthemodernfauin theIsthmianregionthat na is withSouthAmerica.This was recognizedearMa a sill beganforming
flowof the GulfStream.This provided ly on by Wallace (1876): "The portionof North
intensified
warmwaterto the NorthAtlanticregionthatto- Americathatlies withinthetropics(MexicanSubgetherwith Milankovitchorbitalcycles, and de- region)closelyresemblesthe [BrazilianSubregion]
fromwan- in generalzoologicalfeatures,"and he noted"unCO2 concentration
creasingatmospheric
ing plate tectonic activity,eventuallyproduced mistakableevidence of an extensiveimmigration
of glacial fromSouthintoNorthAmerica,notlongbeforethe
conditionsfavorableto the development
Glacial epoch" (quotedfromWebb& Rancy,1996:
climates.

558

Annals of the
Missouri Botanical Garden

depths
Abysoal-bathyal

Figure11. CentralAmericanIsthmusduringthe middleMiocene(16-15 Ma). Oblique parallellines = emergent

land; dots = shelfsediments;parallellines = abyssaloceanic sediments.FromCoatesand Obando(1996). Used with
of ChicagoPress.
permissionofthe University

fromnorthto
335-336). Second, large numbersof immigrants The flowofmammalianimmigrants
fromNorthand CentralAmericaare firstfoundin southappears to have been largerthanfromsouth
SouthAmericaaftertheestablishment
oftheland- to north(Marshallet al., 1982; Webb, 1985). In
bridge.Relativelyfewer,but still a large number, addition,thereappearsto have been a higherspeof SouthAmericanmammalgeneraare firstnoted ciationrate amongtaxa thatmovedto the south,
in Northand CentralAmericaafterthelandbridge compared to the northward-moving
immigrants
numbersof
was completed.These Pliocene migrantswere (Fig. 14). Thus, in additionto different
oftemperate
mostlycharacteristic
climatesand sa- species successfullymigratingin each direction,
vanna habitats.Evidence fromfossil mammalian therewerealso different
ratesofsubsequentdiverfaunasis further
to suggestthatby the sificationamongmammaliantaxa involvedin the
interpreted
middlePleistocenethesavannacorridor
patThis resultedin theasymmetrical
was giving interchange.
way to tropicalrainforest,
amongextantNorthand South
especiallyduringinter- ternof distribution
glacial times(Webb& Rancy,1996: 348).
Americanmammals.Accordingto Webb (1991),

Figure12. CentralAmericanIsthmusduringthelate Miocene(7-6 Ma). Symbolsand sourceas in Figure11.

Volume 86, Number2

1999

Burnham& Graham
Historyof NeotropicalVegetation

TempFgqur-S1n

559

Carloc-

Figure13. CentralAmericanIsthmusduringthelate Pliocene(-3 Ma). Symbolsand sourceas in Figure11.

roughly
53% oftheextantSouthAmericanmammal America,near the end ofthe MioceneEpoch, was
fauna is derivedfromdescendantsof immigrants a raccoonknownfromthe late Mioceneof ArgenfromNorthAmerica,whileonly10% ofthe extant tina(Webb,1985), whiletwogeneraofSouthAmermammalianfaunaof NorthAmericais ascribedto ican groundslothsappear in the late Miocene of
descendantsofthe interchange.
NorthAmerica.By the late Pliocene and PleistoorAn earlyintroduction
fromthe northintoSouth cene, the numberofland mammalsofnorthern
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560

Annals of the
MissouriBotanical Garden

iginin SouthAmericaincreasesto 15 familiesand et al., 1997), while manyof those fromnorthern

29 genera (Webb, 1985: table I). These include SouthAmericaare fromthehighlandsofColombia
modem,extinct,or ancestralgeneraofthe Chirop- (Hooghiemstra,1984; Hooghiemstra& van der
tera (bats), Soricidae (shrews),Leporidae (hares), Hammen,1993; van derHammen& Hooghiemstra,
Heteromyidae(pocket mice), Geomyidae(pocket 1995; 2000-4000 m). However,studiesfromupet
gophers),Sciuridae (squirrels),Sigmodontinero- land depositsin CentralAmerica(Hooghiemstra
of Carniv- al., 1992, Iselbe et al., 1996, Costa Rica, 2310 m;
dents(fieldmice),the firstintroduction
ora-Felidae [cats; Smilodon(New WorldPleisto- Martin,1961, Costa Rica, 2400 m) and fromthe
cene saber-toothed
tiger),Puma, Panthera,Felis], lowlandsof Amazonia(Colinvauxet al., 1996a, b;
Mustelidae(weasels, skunks,otters),Canidae (fox Liu & Colinvaux,1988; van der Hammen& Absy,
and wolf),Procyonidae(raccoons),Ursidae(bears), 1994) allow new calculationsfromcomparableloreGomphotheriidae
(elephantids),Tapiridae(tapirs), calities. Appendix2 lists 235 palynomorphs
Equidae (horses),Tayassuidae(peccaries,pig-like portedfromQuaternarydepositson both sides of
forms),
earlyCamelidae(camel line),and Cervidae the Isthmusof Panama. Seventyof the palynomorphsoccur both in southernCentralAmerica
(deer).
SouthAmericafora similarity
valGroupsmovingfromSouthAmericaintoCentral and in northern
and NorthAmericainclude the Dasypodidae(ar- ue of29.7%. Thus,thepaleobotanicaltrendis conrecord,
madillos), Mylodontidae(sloths), Glyptodontidae sistentwiththatof the marineinvertebrate
(an extinctgrouprelatedto thearmadillos),Ereth- tectonics,and mammalianfossilfaunas.The floras
(through
about
izontidae(porcupines),Hydrochoeridae
(capybara), are distinctuntillate in theTertiary
Trichechidae(manatees),Phorusrhachidae(large the Mio-Plioceneinterval)withgreatersimilarities
predaceous ground birds), Megatheriidae(giant developingin the Plio-Pleistoceneand especially
groundsloths),Megalonychidae
(largeextincteden- in the Pleistocene.
tate mammals),Camelidae (represented
by llamalike camelids),Didelphidae(opossums),and Toxo- Habitatsoftheinterchange
dontidae(rhino-like
ungulates).
The fossilplantdata are also generally
consistent
The mammalianrecordindicatesthat connectionssuitableforthe migration
ofdry-landspecies withestimatesof habitats(savannas) and climate
based on mammalianfossilfaunas,but
across the Isthmuswere essentiallycompleteby (temperate)
The paleobotanicalevidence
-2.5 Ma. Accordingto Webb and Rancy (1996), withsomedifferences.
tropicalforestthroughout
savanna habitatswere at least locallyavailable at indicatespredominantly
grassysavannas dethattime.The originof the modemtropicalmam- the Cenozoic,withrestricted
malianfaunaof CentralAmericais placed at ap- velopingin aboutthelate Miocene.The earlyMiocene microfossil
florasfromPanama (Culebra,Cuthe middlePleistoceneor -780 ka.
proximately
The Cenozoicfossilplantrecordalso reflects
dis- caracha, La Boca; -90N latitude;Fig. 15) and
tinctSouthAmericanand CentralAmericanvege- Costa Rica (Uscari; -10'N latitude)containvirtuAs ally no pollenofthe Gramineae,whilein theMiotationthroughthe early and middle Tertiary.
notedpreviously
here(Table 1), in theearlytomid- PlioceneGatunfloraofPanama,grasspollenreachdle Eocene only-2.6% ofpalynomorphs
are com- es a maximumof 7.5%. This marksthe beginning
monto assemblageson bothsides of the Isthmus. of at least local savannasthatpresumablycontinthePliocene,and expanded
In the earlyMiocene the figureis 10.7% and de- ued to developthrough
as reflected
bytheincrease
creases to 8.9% in the middlePliocene. The de- duringglacialintervals,
crease correlateswithrisingsea levels duringa in grazingand browsingmembersof mammalian
middlePlioceneintervalofincreasingwarmth
(Cro- faunas.However,in theCenozoicpollenflorasfrom
nin & Dowsett,1991). The variousfactorsthataf- southernCentralAmerica,the dominantcompoare tropicalforestelfect the calculations are discussed by Graham nentsof the paleovegetation
betweenmam(1992), and it is especiallyimportant
to notethat ements.This apparentinconsistency
in palynomorphs
similarities
and not malianfaunasindicatingopen savannavegetation,
theyrepresent
in vegetationdirectly.In the originaltabulations, and paleobotanicalevidenceformostlyclosed tropthe percent similarityfor the late Quaternary ical forest,is paralleled in the Tertiaryrecordof
NorthAmerica.Theretheexplanation
because most northwestern
(15.7%) was especiallyapproximate
ofthevegetation
disturbance
of the fewstudiesavailable fromCentralAmerica appearstobe frequent
werefromthelowlands(e.g.,Bartlett
& Barghoorn, by ash falls associated withextensivevolcanism
1973, Panama; Bush & Colinvaux,1990, Panama; (Cross & Taggart,1982; Taggart& Cross, 1990;
as a
Costa Rica; see also Phillips Taggartet al., 1982). This was interpreted
Horn,1985, off-shore

Volume86, Number2
1999

Burnham
& Graham
ofNeotropical
History
Vegetation

561

SIMOJOVEL

SOLO

LAKE VALENCIA

CUCARACHA
CULEBRA

PETEN

GATUNCILLO
LA BOCA
PADREMIGUEL/
HERRERRIA
0,

EVLE
PAA

~~~~~~~~LAYEGUADA
CAQUETA

<-ER
J

SAN JUAN BOSCO

Figure 15.

HASCARAN

LRONDONIA

CARAJAs-

SALITRE

General location of fossil palynoflorasmentioned in the text.

shifting
mosaic of open, short-lived
habitatssuffi- Period.Throughthe earlyMioceneall elementsin
cientto supportmobileherdsofbrowsing
and graz- thefossilflorasfromsouthernCentralAmericacan
ing ungulateswithina mostlytemperateforested be accommodatedat elevationsbetweensea level
vegetation.The frequentash layersin the Cucara- and -1400 m. By the Mio-Pliocene(Gatunflora)
cha Formation
ofPanama suggestsa similarmech- paleoelevations,based on modernanalog distribuanism(Graham,1988b). Evidenceofextensivesa- tions,had increasedto -1700 m (lowermontane
premontane
vanna developmentis onlypresentin Quaternary moistforest,lowermontanewetforest,
floras,but is neverabundantin CentralAmerica. dryforest,montanemoistforest,montanewetforelevationsare 3820 m (CerComparabledata have not yet been presentedin est). Presentmaximum
South America.Savanna, as recognizedtodayin ro Chirrip6)and 3432 m (VolcalnIrazui)in Costa
CentralAmerica,may be a Pleistocenedevelop- Rica, and 3475 m (VolcalnBari) in Panama. The
mentaugmentedby recentanthropogenic
factors. appearanceofthesehighlandsin theMio-Pliocene
winds fromthe
Thereis also evidenceto suggestthatdrierhab- began to deflectmoisture-laden
betweena
itatssupporting
moreopen vegetation
begandevel- northand initiatedthe differentiation
oping locally in the late Miocene and earlyPlio- wetterAtlanticand drierPacificside,as at present.
cene. The early Tertiaryfloras from southern
The development
oftemperate
habitatsin protoCentralAmericacontaina fewelementsofdryhab- CentralAmericaduringthe Middle and Late Ceitats.By the Mio-Pliocenetheseelementsincrease nozoicis complex.The initialappearanceofnorthand include 11 taxa that collectivelysuggestan ern temperateelements is in the Mio-Pliocene
earlyformof tropicaldryforest-Poaceae,Acacia, Gatunfloraof Panama,wherea fewpollen grains
cf. of Quercusare firstrecovered.This correspondsin
Allophylus,
Busera,Cedrela,Ceiba, Combretum,
Jatropha,
Posoqueria,Pseudobombax,
and Serjania timeto the appearanceof the paleoelevationsthat
theregionintowetterand dri(Graham,1991).
begandifferentiating
It is likelythatthese local drierconditionsre- er sides. By the late Quaternary
pollen of Alnus,
sulted,at least in part,froma developingrainshad- Juglans,Liquidambar,Myrica,Quercus,Salix, and
ow due to increasingelevationsduringtheTertiary Ulmusis presentin terrestrial
CentralAmerican

562

Annals of the
MissouriBotanical Garden

Table 3. Laurasian genera now in tropical South

sediments,althoughJuglansand Liquidambarin
off-shore
depositsfromCosta Rica (Horn, 1985) America.SouthAmericanspecies numbersare approxifromfarther
north.Lo- matedfromguides and floras(e.g., Gentry,1993; Brako
mayhave been transported
calized temperatehabitatsfirstappear in the Mio- & Zarucchi,1993; Killeen et al., 1993).
Pliocene and it is likelytheyresulted,at least in
Shrubsand herbs
Trees
part,fromincreasingelevationsratherthanexcluAmbrosia(?)
Alnus
(1)
sivelyfrommarkedchangesin climate.We assume
Berberis(32)
Boehmeria(8)
thatthe low-lying,
peninsulararea in questionreErigeron(?)
Cornus(1)
ceived ameliorating
sea breezes and precipitation
Lupinus(171)
Morus(2)
that may have dampened the global signal of
Ribes(16)
Myrica(3)
and
changescaused bytrendsin CO2concentration
Rubus(18)
Prunus(19)
solarfluctuations.
Salvia (76)
Quercus(1)
climates,especially
Anotherfactorinfluencing
Rhamnus(2)
Satureja(26)
along coastal areas, is upwelling.As the Panama
Scutellaria(15)
Salix (2-3)
Sambucus(1)
Valeriana(100)
landbridgebecameestablished,theflowoftheGulf
(9)
Vaccinium
and broughtcold bottomwater
Streamintensified
Viburnum
(12)
to the surfacein someregions.This is a likelyexplanationforthe cooler climatesindicatedby the
middlePliocene Paraje Solo floraof southeastern
Veracruz,Mexico(17'N), duringan intervalofover- Simpson (1975) documented taxa that,based on exall warming(Graham,1976, 1998b). Ocean circu- tant distribution, must have been immigrants to
(van der South America fromthe north.The list is short,and
lation,orogeny,
lowerCO2 concentrations
et al., 1996), concen- few genera show radiation followingtheir arrival in
Berget al., 1993; Ktirschner
trationsfromgloballywaningpost-Mesozoicplate South America. Woody plant taxa that are well
in known for their appearance in South America foltectonicactivity,
and solar-inducedfluctuations
climate via Milankovitchmechanismsall influ- lowing establishment of the landbridge include AlIn pro- nus, Myrica, Quercus, Cornus, and Salix (Table 3).
encedtheCenozoichistory
ofenvironments.
to-CentralAmericaa combinationof factorsperi- These genera include only one or a few species in
odicallyproducedlocal conditionssuitableforthe South America (Brako & Zarucchi, 1993), although
oftemperate
biotasofopen habitatsin theycan be ecologically importantin many Andean
interchange
thelate Tertiary
Period.These conditions
prevailed areas. Pollen records cannot be used to resolve the
especiallyin thehighlandsand becamemoreprev- issue of speciation in these cases because the polPeriod.
len cannot be recognized to species. A more evoalenttowardthe end of theTertiary

Comparison
of themammalianand angiosperm
records
and speComparisonofthepatternofmigration
ciationbetweenplantand animalgroupsinvolved
in theexchangeacrossthePanamanianlandbridge
littleattention.
has receivedsurprisingly
Simpson
beand Neff(1985) noted the lack of similarity
tweenplant and mammalrecordsbut did not expand on the specificpatternsin each lineage.The
plantdata requiredforsuch a comparisonis based
on two sources of information.
First,directfossil
evidencefromcomparisonoftaxonomiclistsoffossil pollen in Colombiaand the Guianas withfossil
pollen in Panama and Costa Rica (Graham,1992)
as well as comparisonof sites offirstfossiloccurrences of taxa believed to have been involvedin
the interchange(Taylor,1988), and second, evidencefrommoderndistributions
ofplanttaxa(Gentry,1982a, 1992; Hammel& Zamora,1990, 1993;
Wendt,1993). Raven and Axelrod (1974) and

lutionarilysuccessful group of shrubs and herbs includes Berberis, Lupinus, Rubus, Scutellaria, and
Viburnum(Table 3). The South American species
of these genera verylikely came into existence after
their immigrantancestors reached South America,
similar to the radiation proposed withinmammalian
genera. However, the timing of the immigrantarrival, relative to the establishment of the landbridge, is difficultto establish withoutdirect information from the fossil record. Their arrival could
predate the landbridge, and in this case examination of dispersal mechanisms might prove useful.
These plant taxa are absent or are recovered rarely
fromthe palynological record, most likely because
they are insect-pollinated. The arrival of the successful sigmodontinerodents was proposed to have
predated the landbridge (Hershkovitz, 1966; Reig,
1980, 1986; Marshall & Sempere, 1993), but a consensus has not been reached on this point among
vertebrate paleontologists (Patterson & Pascual,
1972; Flynn et al., 1985; Marshall & Sempere,
1993). The fact that South American sigmodontines

Volume 86, Number2

1999

.Burnham

& Graham
Historyof NeotropicalVegetation

563

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::::::::.,.,::.:.:.:.:.::::::::::::::::::::::::::::::::......:
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thanNorthAmerican R. Anderson, pers. comm. 1997). These include raappear to be moreprimitive

and in host- diations into the moist forests of Mexico (Wendt,
forms,at least in penile morphology
casts doubtson theirarrival 1993), but appear to be more typical of the dry
parasiterelationships,
in South Americavia the landbridge(Jacobs & tropical forests of Central America, which were
pers.comm.1998). widespread until recent deforestation (Janzen,
Lindsay,1984; B. D. Patterson,
patternviewedfromSouth 1988).
The plantimmigration
In summary,the plant and animal migrationand
ofthelandbridgereAmericais thatestablishment
of radiation patterns are interestinglydifferent(Figs.
changesin the composition
sultedin significant
SouthAmericanuplandforests.However,thenum- 14 and 16). Mammalian radiations in South Amerber ofspecies,and certainlythe numberofgenera ica are noted in many differentgroups of larger
is low. The mammals (deer, canids, camelids, tapirs), but are
immigrants,
descendedfromnorthern
less
significant particularly importantamong the sigmodontinerofar
been
landbridgeappearsto have
for speciationin lowland angiospermsin South dents. These rodentsare abundant in manyhabitats
Americathan it was formammals.To our knowl- throughoutSouth America, and are perhaps most
edge,thereare no estimatesofthe numberofspe- species-rich in uplands and open habitats (Hershcies in the SouthAmericanflorathatwerederived kovitz, 1972). However, when their current distributwe wouldplace it at bution is mapped on their phylogenetic diversifiimmigrants,
fromnorthern
cation it appears that lowland sigmodontines gave
about 10%.
Wendt(1993) doc- rise to highland sigmodontines, rather than being
perspective,
Froma northern
that derived fromthem (Patterson, in press). This conumentedtaxa based on moderndistributions
fromSouth America trastswith the situation in plants where uplands do
had most likely immigrated
and speciated in Mexico and Central America. correspond to the site of ecological dominance of
These were not comparedto the mammalianpat- the immigrantangiosperms from North America,
terns.RadiationsintocentralMexicoand the An- but in the case of immigrantwoody plants, ecologtilles have been suggestedby specialistson Big- ical dominance is not accompanied by evolutionary
noniaceae, Malpighiaceae, and Sapindaceae diversification.Once arrivingin Central and North
1993, 1996; W. America, southernmammals were relativelyunsuc1992; Acevedo-Rodrfguez,
(Gentry,

564

Annals of the
Missouri Botanical Garden

thereare no fossilflorasof Paleocene age known

eleLatinAmerica,north-temperate
fromnorthern
mentsare absent fromthe Eocene Burgas Basin
assemblage in northernMexico (Graham,1999a;
et al., 1980), the Gatuncillo
Martinez-Hernandez
assemblagein Panama (Graham,1985), and from
earlyMioceneassemblagesin CostaRica (Graham,
1987) and Panama (Graham, 1988a, b, 1989).
These elementsare knownfroma few grainsof
Quercusin the Mio-PlioceneGatun Formationof
Panama,and froma richerassemblagein themidofsoutheastern
dle PlioceneParaje Solo Formation
Veracruz,Mexico(Abies,Picea, Pinus,Alnus,Celtis,
Juglans,Liquidambar,Myrica,Populus, Quercus,
and Ulmus).Based on thisevidence,itappearsthat
temperatecomponentof
the originof the northern
the Latin American vegetation was primarily
If norththroughrelativelyrecent introductions.
elementin
Originof thenorthern
temperate
temperateelementswere presentpriorto the late
neotropical
vegetation
Miocene,thesephantomforestsleftno substantive
Twoscenarioshave been proposedfortheorigin recordin thefossilassemblagespublishedto date.
ofnorth-temperate
plantsin Mexico,CentralAmerUntil recentlythe numberof fossil florasby
ica, and northern
SouthAmerica.One is thatmany whichthemodelscould be evaluatedwas relatively
of these elementswere part of an ancientwide- low. New data frommicrofossil
florasin Chiapas
LatinAmer- (Mexico),Guatemala,and Costa Rica providefurspreadvegetationexistingin northern
ica since thePaleogene(Axelrod,1975). The other therinformation
on the principaltimeoforiginfor
is thatmanymigratedsouthward
withcoolingcli- north-temperate
LatinAmerelementsin northern
mates,the appearanceof upland habitats,and the ica. The evidencepreviouslydiscussedwouldsugtheIsth- gest thatfossilflorasolder thanabout the middle
establishment
ofland connections
through
mianregion(Graham,1973). Thereis also thepos- to late Mioceneshouldcontainfewor no represensibilityof random and occasional introductions tativesof these elements;rather,theyshould be
transport. presentin florasofyoungerage, and theirdiversity
throughout
theCenozoicbylong-distance
Three tests are available forassessing the likeli- in the fossil florasshould decrease towardthe
hood of each model. First,if therewas a general south.The Simojovelflorain Chiapas, Mexico,is
fromthe northintoMexi- EarlyMiocenein age (Frost& Langenheim,
1974),
progressiveintroduction
ofdeSouthAmerica, and it is withinthe present-day
distribution
co, CentralAmerica,and northern
the oldesttemperateplantfossilsshouldbe in the ciduous trees and shrubs disjunct fromeastern
northwiththeyoungerones towardthesouth.Sec- NorthAmerica (Breedlove,1973; Gomez-Pompa,
intonorthern
SouthAmerica 1973; Miranda& Sharp,1950). Withtheexception
ond,theintroductions
shouldcoincidegenerallywithclosureoftheIsth- ofPinus,thefossilfloracontainsno representatives
mus; thatis, theyshould mostlyappear in South of easterndeciduoustemperateelements(Graham,
Americaafterabout the Mio-Pliocene.Third,the 1998a, 1999b; Graham& Palacios Chavez, 1996;
introductions
into northern
Latin Americashould Langenheimet al., 1967). Temperatehabitatswere
withsomeglobalclimaticeventthatfa- available, but theywere occupied by plantssuch
correspond
of north-temperateas Podocarpusand Engelhardia.
voredthe southwardmigration
In Guatemala,a late Miocene to Mio-Pliocene
organisms.
In the southeastern
UnitedStates,pollensimilar assemblageof pollen and spores has been recovto Alnus,Betula, and Carya is presentin deposits ered fromthe Padre Miguel-Herrerfa
complexof
of Paleocene age (Frederiksen,1991). In the Eo- formations
(Graham,1998a). Picea, Pinus,Juglans,
cene (Claibornian)thereare added Pinus, Casta- Quercus,and Ulmushavebeen identified,
revealing
nea, Celtis,Fagus, Quercus,Nyssa,Juglans,Os- a north-temperate
componentin the late Miocene
Platanus, Tilia, and Ulmus,and in and Mio-Pliocenethatwas absentin theolderearly
trya-Carpinus,
the youngerJacksonianPicea, Tsuga,Acer(?),Di- Miocene Simojovel florafromadjacent Chiapas.
ospyros(?),
Fraxinus(?),and Nyssa appear (Freder- Farthersouthin Costa Rica theseelementsare,abiksen, 1981, 1988, 1991; Gray,1960). Although sent in the Pliocene Rio Banano Formation(Gracessfulin an evolutionary
sense,whereonlyone or
a fewspecies ofeach group(sloths,capybara,agoutis,and opossums)are found(Emmons,1990). Conplants exversely,while some northward-moving
tendjust intothewetforestsofPanamaor southern
Costa Rica, the moresuccessfulplantgroupsthat
northshowstrongdiversification
into
extendfarther
thedrier,seasonalhabitatsofCentralAmerica.The
topollen recordsthat are rapidlyaccumulating,
of species lists fromboth
getherwithrefinements
sides ofthe landbridge,shouldmake it possibleto
establish the ecological conditionsunder which
Withthisadded ecologicalperspectaxa migrated.
tiveit maybe easier to postulatereasonsfora parin
ticular migrationevent and the environments
whichspeciationmayhave occurred.

565

Burnham& Graham
Historyof NeotropicalVegetation

Volume 86, Number2

1999

TEMPERATE ELEMENTS
RARETOABSENT j PRESENT
BURGOS BASIN

_16
12-

MENDEZ-?

12
;

CULEBRA IXTAPA

<

SIMOJOVEL

PADRE MIGUEL/
HERRERIA- 5 GENERA

PINUS

RIO BANANO - ILEX

-8
4-I

PARAJESOLO 10GENERA

NONGLACIAL
GLACIAL
-4

0-

_00
E
_
FCREEMODERN
TC

70Ma

60

50

40

30

PLIO
5.2 1.6

MIOCENE

OLIGOCENE
35
23

EOCENE

PALEOCENE
65
56
I

20

10

Figure 17. Benthic paleotemperature curve, TertiarypalynoflorasfromnorthernLatin America mentioned in the

text, and occurrences of northerntemperate elements. Paleotemperature curve based on Miller et al. (1987).

ham & Dilcher,1998). Thus, northern

temperate species (Brako & Zarucchi,1993; Killeen et al.,
elementsare absentfromtheEocene BurgasBasin 1993).
Flora of Mexico, the Eocene Gatuncillofloraof
The thirdtest is whethersome climaticevent
the
timethatfacilitated
Panama,theearlyMioceneSimojovelfloraofChia- occurredat theappropriate
elementsintoLatin
ofnorth-temperate
pas, Mexico,the earlyMiocene Uscari Formation introduction
and the Pliocene Rio Banano Formationof Costa America,and thatis evidentfroman independent
Rica, and the earlyMiocene Culebra,Cucaracha, line of inquiry.In Figure17 the florasare plotted
curve. A
and La Boca florasofPanama.Theyare presentin on the global benthicpaleotemperature
is evidentin themiddle
Mio-Plioceneflorasof Guatemala;in the middle majordropin temperature
PlioceneParaje Solo floraofVeracruz,Mexico;and Miocene that initiatedAntarcticglaciations,exand conglacierson Antarctica,
just appear (Quercusonly)in theMio-PlioceneGa- pandedcontinental
tunfloraofPanama.
tributedto the spreadof seasonallydryhabitatsin
The second testis whetherthesetemperate
ele- NorthAmerica(see discussionin Graham,1999c,
is conmentsappear in northern
SouthAmericaprimarily chapters3 and 7). This dropin temperature
based
afterclosureoftheIsthmusofPanama (afterabout sistentwiththe timingof the introductions
the Mio-Plioceneinterval).In SouthAmericaSalix on the available paleobotanicalevidence,and proappearsin theearlyPliocene,Myricain themiddle vides a plausible mechanismforthe earlyappearPliocene,Alnusat - 1 Ma, and Quercusat 330 ka ance of the northern
temperatecomponentin the
latEvidencefortheslightly
(Hooghiemstra,
1989, 1994; Hooghiemstra
& Ran, neotropical
vegetation.
& Sarmiento,1991; Wijinga, er appearance of uplands (Mio-Pliocene)supple1994; Hooghiemstra
1996). One exceptionmaybe seeds ofJuglansre- mentingeffectsofLate Cenozoiccooling,has been
portedfrompresumedMiocenedepositsin Ecuador discussedpreviously(thispaper).The pre-existing
(Brown,1946), but Manchester(pers.comm.1997) uplands and risingmountainareas in Centraland
datesare neededtocon- South Americawould have been expectedto exsuggestedthatradiometric
firmtheage ofthespecimens.Juglansmaybe rep- perience more pronouncedclimate changes than
resentedin SouthAmericatodaybyas manyas five the lowland,coastal areas. It is theseuplandareas

566

Annals of the
Missouri Botanical Garden

group,tribeRobinieae and
thatlikelyprovidedhabitatsfortheearlytemperate groups(Dichrostachys
elements.
allies; Lavin & Luckow,1993). If a taxonhas a
and an earlyTertiary
modemcenterofdistribution
in
North
America,thenthere
tropical
fossil
record
Originof thetropicalelementin thenorthern
willbe in
thatits sister-group
is a highprobability
LatinAmericanvegetation
the paleotropicsand derivedrelativeswill be in
New analysesalso are providing
insightsintothe SouthAmericaif it has been introducedfromthe
Latin northfromGondwananancestors.The resultsofthe
originof the tropicalcomponentin northern
Americanvegetation.It has generallybeen as- analysisfortheselegumessupportedthe hypothesumedthattreesand shrubsin thetropicalforests sis; namely,thetaxain SouthAmericawerederived
tropical
of Mexicoand CentralAmericawerederivedfrom from,and not a source for,the northern
SouthAmerica:"Indigenoustropicalplantsin mod- ones. In contrast,a similaranalysisof the Costa
and Rican Ruptiliocarpon
em NorthAmericaare descendantsofTertiary
in Lepidobotryaceae(HamfromSouthAmerica"(Cron- mel & Zamora,1993) showedthatit is likelyan
Quaternary
immigrants
quist,1988: 153). In generalthisis true,but it is exampleof the majorityof taxa thatarrivedfrom
provingtoo simplistic.
South America.The emergingpictureis thatthe
Wendt(1993) studiedthe affinities
of rainforest tropicalforestsofnorthern
LatinAmerica,although
canopytreesin Mexico.As expected,the majority primarily
SouthAmericanin origin,are morecomof the taxa (75%) are related to ones in South plex thanearlierthoughtand are composedof elAmerica,and probablyarrivedvia an Isthmian ementsarrivingfromseveral sourcesvia different
of the Panama landbridge, routes and at differenttimes (Graham, 1995;
route afterformation
and/orby long-distancetransportthroughout
the Wendt,1993).
Cenozoic Era. However,the modem distribution
and affinities,
and the fossilrecordofa significant EVENT IV: QUATERNARY CLIMATE FLUCTUATIONS
number(-25%), suggesta different
history.Acand
The firstbroadassessmentof global Quaternary
cordingto Wendt(1993), moderndistributions
relationshipsindicatethatsome are of Laurasian climateswas fromthe ClimateLong-RangeInvesfromthenorthvia the tigationand MappingProgram(CLIMAP) project
originand probablymigrated
NorthAtlanticlandbridge-Antirhea,
Aphananthe, (CLIMAP Project Members,1976, 1981, 1984).
Berrya,Bourreria,
Bursera,Ehretia,Erblichia,Fed- The resultingmodel,based on changesin thecomofplanktonicmicrofossils
erovia, Flacourtiaceae (subtribeHydnocarpinae), positionand distribution
indicatedthatwhile highlatiand Tri- (e.g., foraminifera),
Lonchocarpus,
Sideroxylon,
Styphnolobium,
The fossil recordprovidesevidence tudeschilledat glacial maximumat - 18 ka, temchospermum.
forothersofLaurasianoriginthatfolloweda north- peraturesin the low latitudesremainedabout the
ern routeintoMexico.These are taxa knownfrom same or to a maximumof about20 cooler.This is
the Eocene of the southeastern
UnitedStates,but at variance with new oxygen isotope evidence
whichpresently
have no or onlylaterfossilrecords (Guildersonet al., 1994; Schrag,et al., 1996), noin South America-Celtis, Clethra,Cordia, flex, ble gasses dissolvedin '4C-datedgroundwater(StuPopulus, Saurauia, Salix, Talauma, Trema,and te et al., 1995), snow-linedepression(Rind & Peclimaterecordsfrompollen
Turpinia.
teet,1985), continental
A groupof African-Gondwanan
originprobably analysis,and withglobal circulationmodel(GCM)
arrivingfromthe northincludesAcacia, Beilsch- simulationsthatpredicta MAT (meanannualtemmiedia, Bursera, Caesalpinia, Cassia, Cedrela, perature)coolingin the equatorialregionsof 5?Chrysophyllum,
Dalbergia, Diospyros,Ficus, Lon- 6?C at glacial maximum(Colinvauxet al., 1996a,
chocarpus,Nectandra,Ocotea, Oreopanax,Persea, b, and see reviewin Kerr,1997). New calciumratiosfromsubmergedcoral reefs(AcroSabal, Sapindus,Sapium,Sterculia,and Termina- strontium
prior pora palmata) drilledoffthe southcoast of Barlia, whileothersofGondwananoriginarriving
to the landbridgebut stillvia SouthAmericamay bados (13'N) also favor cooler climates. The
includeAlchornea,Allophyllus,
is chemicallysimilarto calcium
Apeiba,Astronium, elementstrontium
Bauhinia, Bernoullia,Casearia, Dendropanax,Er- and can replacecalciumin thewalls ofthecorals.
ythrina,
Eugenia,Homalium,Luehea,Mabea, Och- The rate of replacementis temperature-sensitive,
indicatetemperatures
lowerby
and measurements
roma,Pouteria,Tapiria,Thouinia,and Trichilia.
The presenceofplantsofOld Worldoriginin the - 50C at - 18 ka (Guildersonet al., 1994). ModthemechLatinAmericaalso has elingbyWebbet al. (1997) demonstrated
tropicalforestsofnorthern
been assessed by a cladisticanalysisoftwolegume anismpotentially
responsibleforequatorialcooling.

Volume 86, Number2

1999

Burnham& Graham
Historyof NeotropicalVegetation

567

Otherrecentdata also suggestthatthemiddleand

low latitudesrespondedto fluctuations
in Quaternary temperatures(Charles, 1997; Beck et al.,
2~~~~~~
1997; Curry& Oppo, 1997; Webbet al., 1997). A
cool intervalcalled the LittleIce Age occurredat
- 400 yearsago and at thattimeMAT in the Sargasso Sea (- 320N)was - 1C coolerthanat present.Duringa warmintervalat - 1 ka itwas - 10C
warmer(Keigwin,1996).
AmongtheunsettledaspectsofQuaternary
veg100
etationalhistoryin SouthAmericaare the history
of tropicallowlandbiotas and the relatedHaffer
modelof diversification.
Earlypollenprofilesfrom
theHighPlain ofBogota'revealedthattreelinefluctuatedby - 1700 m duringthe late Pliocene and
Quaternary(van der Hammen& Gonzalez,1960,
1964). This corresponds
in MAT of
to a difference
- 80C at - 2500 m; thepresentMAT is 17-19'C.
0~~~~
By extrapolation,
a potentialloweringof up to 60C was indicatedforthelowlandsduringthecoldest intervals.Glacial climateswerealso frequently
drierbecause coolertemperatures
reducethe rate
ofevaporationofwaterfromthe ocean surfaceinto
In thenorthern
theatmosphere.
Andes,rainfallwas
periodicallyreduced by up to 50% (van Geel &
van der Hammen,1973). The questionis whether
these changessuggestedby geochemicaland oxygen isotopedata, and evidentin theAndeanhighOSavanna
I* TropicalForest
lands,affected
biotasin thelowlandAmazonBasin.
in AmaFigure 18. Projectedrainforest
distribution
Haffer(1969, 1970, 1974, 1982) observedthat
zonia duringthe Pleistocene,with(A) 25% and (B) 40%
species diversityamongtoucansand jacanas was reductionsin rainfallfrompresent-dayvalues. Shaded
not uniformacross the basin, but was focusedat area showsrainfallover1500 mmannually.Palynological
certainlocales. These areas of highspecies diver- samples fromAmazoniashownby symbols,withinterto sites thatby theirphysiogra- pretedvegetationindicatedby symbolcoding.AfterAbsy
sitycorresponded
phyappearedlikelyto perpetuatemoistconditions and van der Hammen,1994; Colinvaux,1996; Behling,
1998. Sites are (informal
names):1, La Yeguada;2, Lake
duringperiods of aridity(rivermargins,base of Valencia;3, Guianas;4, Mera; 5, San JuanBosco; 6, Camountainslopes). The modelenvisionedrainforest queta River;7, Lake Pata; 8, Carajas; 9, Katira(Rondonbiota concentratedin refugiaduringcooler/drier ia); 10, Lagoa Campestre;11, Catas Altas; 12, Lagoa Cuglacial cycles,whiledrierforestand savanna-like ruca.
area. During
vegetationoccupied the intervening
warmer/moister
interglacialcycles, the rainforest
directly
expandedfromthesesitesand coalesced,whilethe Lacking was paleobotanicalinformation
driercommunities
eitherpersistedon edaphically fromthe lowlandAmazonBasin, and ancillaryinon neotropicalQuaternary
environments
drytablelandswithsandstonesubstratain thelow- formation
lands, or occupied the surrounding
slopes, as at derived fromindependentlines of inquiry.New
and reuniting data were recentlypublished for lower-elevation
present.The periodicfragmentation
ofrangesfacilitatedspeciationbyallopatryand hy- sites aroundthe AmazonBasin and forotherlobridization,and was proposedas one mechanism cationsin the lowlands(Fig. 18). Consideringthe
ofthetropicalecosystem,
and
accountingfor the high diversityof the lowland immensecomplexity
and limitations
inherentin trying
to
tropicalbiota.Severalotherlines ofevidencewere thedifficulties
as supporting
thata coninterpreted
the model,includingthe reconstruct
its past,it is notsurprising
presenceofstonelines whichformunderaridcon- sensushas notyetbeen achieved.Onlya fewplantof plantand insect bearingsites thatextendintothelast glacial interditions,and the concentration
as refugiaon thebasis of val (older than - 12 ka) have been published.
species at sites identified
bird distribution(see papers in Prance, 1982). Haberle (1997) reporteddata fromAmazon fan

568

Annals of the
MissouriBotanical Garden

the lowlandswithout
cores spanning160,000 years thatshow percent- aturesprevailedthroughout
ages of grass and shrubpollen less thanor equal appreciabledryness.One complicationis thaton
as indicatinglimited the basis of meterologicalevidence, the area of
to 25%, a value interpreted
in theAmazonbasin.As not- Lake Pata also has been considereda refugium,
savannadevelopment
ed by Haberle (Haberle, 1997; see also Kerr, and by the Haffermodel it would be expectedto
1996), a few hundredsites in the AmazonBasin supportforestduringdryintervals(van der Hammaybe necessaryto geta clear pictureofenviron- men& Absy,1994). In addition,largereproductive
mentalchangeand bioticresponse.In assessingthe standsofPodocarpusare foundat sea level in Cenit is also necessaryto distin- tral America, on both the Atlanticand Pacific
presentinformation
and Coasts,whichthrowinto questionthe use of this
guishbetweenevidenceforcool temperatures
indicator.
foraridity.There is a broad associationbetween genusas an exclusivelycooler-climate
in the
Data fromothersites suggestthata patchwork
warm-moist
climatesand tropicalvegetation
lowlandsduringthe interglacials(as at present), of vegetationmayhave existedin the lowlandsat
and cool-dryclimatesand drierforest/more
open the coldestintervalswhichincludedmoistforests
savanna-likevegetation
duringglacial times.It has and drier communities.At Carajals(southeastern
been suggested,however,that significant
cooling Amazonia, Brazil; 60S), pollen profilesreflect
occurredin the lowlandsof Amazoniawithoutthe changes in vegetationforthe last 60 ka (Absyet
accompanyingdrynessthatwould have converted al., 1991). The site is on a table mountainat 700areas occupiedbyrainforest
to savanna(Colinvaux, 800 m elevation,today covered withforestand
1996). Thereare insufficient
datafordogmaticcon- edaphic savanna, and surroundedby Amazonian
as
clusionsaboutQuaternary
in theAm- forest.The pollen diagramswere interpreted
environments
azon, and an even-handedreviewmust include showingthatwet intervals(foreston the hills and
in the lowlands;lake levels high)alternatedwith
muchthatis a matterofopinion.
Alongthe easternslopes of the Andes at Mera drierintervals(expansionof savanna on the hills
( 1'S; 1100 m elevation)and San JuanBosco (and in thelowlands;lake levels low).At Rond6nia,
30S; 900 m elevation)in Ecuador,pollen profiles Brazil (90S) in lowlandAmazonia,the sequence is
fromprimarily
to savanna
rainforest
(interglacial),
showthattheupperrangeoftemperature-sensitive
plantpopulationswas lowerby - 700 m between (glacial;42,500 + 2,500 to 18,500 yrB.P.),tomodafter - 11 ka (Absy,1979; Absy&
33,500 and 26,500 yrB.P. (latemiddleGlacial)and ern rainforest
thatelementsfromthe uplands intermingled
with van der Hammen,1976; van der Hammen,1972).
lowlandforestcommunities.
This implies a MAT Geomorphicand pollen data fromthe middleCalowerby - 40C (earlierestimates)to 7-90C (more quetalRiverarea in Colombiaand easternPerualso
to showthatlocal vegetationalterrecent estimates;Bush et al., 1990; Colinvaux, are interpreted
1996; Colinvaux& Liu, 1987; Liu & Colinvaux, nated betweenAmazonianforestand more open
1985) and a moistrainfallregime.However,the caatinga(van der Hammenet al., 1992a, b).
In the "Lagoa Campestre"(Lake) of Salitre in
sites are nearthebase oftheeasternAndeswhere
thepresentannualrainfallis - 5 m. This physical southernBrazil (190S; 970 m elevation),pollendithatwould agramsshowan arid phase between- 50 and 40
settingqualifiesas a potentialrefugium
ka (Ledru et al., 1996). This intervalis characterhave remainedmoistduringdryintervals.
At Lake Pata in northwestern
Brazil (00N; 300 ized by low frequenciesof arborealpollen (0.5m elevation)the pollen recordextendsback to - 3%). Asteraceae (18.5-28%) and Poaceae (6942 ka. Throughout
the profile,tree pollen consti- 70%) are dominantamongthe non-arborealtaxa.
tutes70-90% oftheassemblage,whileherbpollen This pollen assemblageis poor in taxa and india shalremainsbelow 10%. Pollen fromtrees that now cates open,treelessvegetationsurrounding
The
growmostly800-1000 m above thePata site were low pond withno fringeof aquatic vegetation.
present(Podocarpus, 10%; associated Ericaceae, low frequencyof arborealtaxa indicatesthattrees
Raweregrowingonlyat a distance.This suggeststhe
Hedyosmum, Humiria, flex, Melastomataceae,
and this presenceoflandscapesfoundtodayin centraland
panea, Weinmannia) at glacial maximum,
was interpreted
to mean a MAT 5-60C lowerthan northern
wherethe climateis cool and
Argentina,
at present(Colinvauxet al., 1996a, b). The infer- dry(Ledruet al., 1996). Otherareas wherereduced
ence is thatat glacial maximumthe area was for- rainfall(- 10-20%) and dryvegetationreplaces
ested, and was not occupied by savanna.Also, if forest in the Quaternaryinclude the Guianas
the area was sufficiently
moistthroughout
thepast (Wijmstra& van der Hammen,1966; Salgado-La- 42 ka to supportforest,
eventhough5-60C cool- bouriau,1980) and the Galapagos Islands (Colinthatlowertemper- vaux, 1972), as well as East Africa(Livingstone,
er,thissupportsthe contention

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Burnham& Graham
Historyof NeotropicalVegetation

569

1975). In an extensivereviewof the geologyand 1990). Considering

thatmuchoftheAmazonBasin
geomorphologyof South America, Clapperton is near sea level, a changeofthismagnitudemust
through
(1993a: 162) concluded,"The important
message have had some dryingeffecton vegetation
is thatclimaticconditions
appeartohavefluctuated fluctuations
in watertable.
betweenhumidand dry,and as the presentinterThereare a numberofindicationsthatlate Glaglacial climateis humid,it seemsreasonableto as- cial and earlyHoloceneclimatesin theneotropical
sume thatdryintervalscorresponded
to periodsof lowlandsincludedcooler and drierintervals,and
global glaciation."
variouslysupporteda mosaic of moistforestsand
A studyof ice cores fromHuascardn in the drierforeststo savanna-likevegetation.
These are:
north-central
Andes of Peru (- 90S, 6000 m ele- (1) fossilmammalianfaunasin Centraland South
vation)has revealed Quaternaryclimaticfluctua- America(Webb& Rancy,1996), (2) a trendtoward
tionsback 20 ka thatparallelthosefromtheNorth drierconditionsand a morecomplexarrayofcomAtlantic(Thompsonet al., 1995). Oxygenisotope munitiesdevelopingnear the end of the Tertiary
ratiosshowthatbetween20 ka and 14.5 ka tem- Periodin Costa Rica and Panama (Graham,1992);
in Guatransitions
peraturesat thiselevationwere8-12'C coolerthan (3) aridityat interglacial-glacial
at present.This wouldbe consistent
withthe - 60C temala (Leyden, 1984; Leyden et al., 1993); (4)
loweringof temperatures
estimatedfor the low- geomorphicfeaturessuggestingaridityduringthe
lands. In addition,therewas a 200-foldincreasein Quaternaryin the Amazon lowlands (Ab'Saber,
dust and lowernitratelevels in the cores.Thomp- 1982; Clapperton,1993a); (5) markedchangesin
son and co-authors(1995: 47) concluded,"The ex- sea level (GeophysicsStudyCommittee,
1990); and
tremedustinessof the LGS [late glacial stage]ice (6) Quaternary
fromotheradariditydemonstrated
on Huascaran is consistentwith reconstructionsjacent and distantpartsoftheworld(theGuianas,
[Clapperton,1993b; van der Hammen & Absy, Wijmstra& van der Hammen,1966; GalapagosIs1994] forSouthAmericathatindicatea reduction lands, Colinvaux,1972; Lake Valencia,Venezuela,
in atmospherichumidity,
precipitation,
and forest Leyden, 1985; East Africa,Livingstone,1975).
eiand grasscoverduringthe LGS alongwithan en- This evidenceis open to otherinterpretations:
hancementof eolian depositssuch as dune fields therthatthelowerrainfallwas insufficient
to cause
or savanna,or
and deflationbasins [Clapperton,1993a], because forestto be replaced by dry-forest
winds were strongerand surfaceconditionswere thatsuch replacements
werelocalized (Colinvaux,
drier."With referenceto the low nitratelevels, 1996).
The consensusdevelopingis thatlate Tertiary
Thompsonet al. (1995: 47) further
notedthey"may
reducedin and Quaternary
sea level and climaticfluctuations
implythatforestcoverwas significantly
responseto dry conditionsand the expansionof affectedthe lowlandbiotas of the Neotropics,and
grassland,as suggestedby palynologicalstudiesin thatthe magnitudeof the temperature
changewas
- 60C. There is less agreementabout aridityand
Brazil [Clapperton,1993b]."
To thenorth,
thehistory
ofQuaternary
vegetation theclosed forestversusmoreopen forestor savanin lowlandPanama is preservedin sedimentsfrom na-likenatureof the vegetation.Colinvaux(1996)
El Valle (500 m elevation;> 30,700 ? 800 to has emphasizedcooler climates,sustainedmoismuch of
8,350 + 150 yr B.P.) and Lake La Yeguada (650 ture,and forestedvegetationthroughout
This wouldnegateany signifm; 14 ka to the present).These recordshave been the late Quaternary.
mechanismsin accountto indicate lowland representation
of icantrole forrefugia-like
interpreted
taxa currently
foundat elevations- 800 m higher ing forthe high level of diversityin neotropical
(Bush & Colinvaux,1990; Bush et al., 1992). The biotas. In contrast,otherauthorssee evidencefor
in thepollen
MAT is estimatedat 4-60C cooler,but stillmoist, periodsofaridityand open vegetation
and thevegetation
as remaining
forested
throughout profilesand fromotherkinds of data. Revisions
the sections.In the Peten of Guatemala,the tem- may become necessaryin the proposedlocations,
peraturedepressionis estimatedat - 80C cooler extent,and patternsof some forestrefugiaand sain the late Quaternary(Leyden,1984; Leyden et vannas in the Amazonianlowlandsas envisioned
truethat
al., 1993), but Leydenalso emphasizedincreased in the originalHaffermodel.It is further
transitions.
climate-induced
refugialeventsin the Quaternary
aridityat interglacial-glacial
In additionto temperature
and precipitation,
an- Period were not the onlymechanismsinfluencing
otherfactoraffecting
lowlandtropicalbiotasis sea- rainforest
Recently,it has been discovdiversity.
A re- eredthatthemodemdistributions
ofsomelineages
level changeassociatedwithglacial history.
cent estimate is for a lowering of - 121 ? 5 m at of rodentsand marsupialsin the Amazon Basin
notto thepresentbarriersformedby river
glacial maximum(GeophysicsStudy Committee, conform

570

Annals of the
Missouri Botanical Garden

van der Hamnmen.

1991. Mise en 6videncede quatre
drainagesystems,but to ancientridgesthatdivide
phases d'ouverture
de la forkt
dense dans le sud-estde
thebasin intosub-basins(Patton& da Silva, 1998;
l'Amazonieau coursdes 60,000 dernieresann6es:Presummaryby Morell, 1996). These ridges formed miere comparaisonavec d'autres r6gionstropicales.
whenthe Andes wereuplifted- 2-5 Ma and diCompt.Rend. Hebd. S6ances Acad. Sci. 313: 673-678.
videdtherangeofsomespecies intoreproductivelyAcevedo-Rodriguez,P. 1993. Systematicsof Serjania.
(Sapindaceae) PartI: A revisionof Serjania sect. Plaisolatedsubsetsthatare stilldetectablebymtDNA
tycoccus.Mem.New YorkBot. Gard.67: 1-94.
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. 1996. Flora of St. John,U.S. VirginIslands.
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P. A. Riezebos & T. van der Hammen.1993. Absolute
othermechanismgenerating
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quence of the Bogota Area, Colombia. Quatern.Sci.
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Madrean-Tethyan
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Appendix1. Publishedplantfossillocalitiesranginggeographically
fromCosta Rica to Bolivia,and temporally
fromLate Cretaceousto Quaternary
Periods.Localitiesare sortedby country,
and withincountry
by geologicage and
thenorderedalphabeticallyby authorof the work.When severalcountriesare treatedby a single publication,the
in alphabeticalorder.
publicationis listedunderthefirstcountry

Country

Geologicalage

Author(s)

Date of
publication

Bolivia
Bolivia
Bolivia
Bolivia
Bolivia
Bolivia
Bolivia
Bolivia
Bolivia
Bolivia
Br. Guiana/
Suriname
Brazil
Brazil

Pliocene
Pliocene
Mio-Pliocene
Mio-Miocene
Miocene
Miocene
Miocene
Miocene
Miocene
Miocene
Paleocene to Miocene
Quaternary
Quaternary

Brazil
Brazil
Brazil
Brazil
Brazil
Brazil

Quaternary
Quaternary
Quaternary
Quaternary
Quaternary
Quaternary

Brazil

Quaternary

Berry
1939a
Berry
1917
Berry
1940
Berry
1922a
Berry
1922b
Britton
1892
Engelhardt
1888
Engelhardt
1894
Gregory
et al.
1998
Singewald& Berry 1922
Van der Hammen& 1964
Wymstra
Absy
1985
Absy& van der
1976
Hammen
Absyet al.
1991
Behling
1995
Behling
1996
Behling
1998
Behling
1998
Behling& Hooghiem-1998
stra
Colinvauxet al.
1996a

Brazil
Brazil
Brazil
Brazil

Quaternary
Quaternary
Quaternary
Holocene

1992
DeOliveira
Duarte& Nogueira 1985
Duarte& Vasconcelos1980
Ledru
1992

Brazil

Quaternary

Brazil
Brazil
Brazil
Brazil
Brazil
Brazil
Brazil

Pliocene
Pliocene
Pliocene
Pliocene
Pliocene
Pliocene
Pliocene

Salgado-Labouriau
et
al.
Berry
Berry
Bonnet
Dolianiti
Dolianiti
Hollick& Berry
Krasser

1935a
1937d
1905
1949
1950
1924
1903

Brazil
Brazil

Pliocene?
Miocene

Selling
De Lima & Angulo

1945
1990

Brazil
Brazil
Brazil
Brazil
Brazil
Brazil
Brazil

Miocene?
Miocene
Miocene
Oligocene
Eocene-Oligocene
Eocene-Oligocene
Oligo-Eocene

Duarte
1972
Hoorn
1993
1992
Mandarim-de-Lacerda
De Lima et al.
1985
Martins-Neto
1989
Beurlen& Sornmer 1954
De Lima & Salard- 1981
Cheboldaeff

1997

Local place name

Organsstudied

Potosi
Potosi& Corocoro
Agua Salada
Pisllypampa
Jancocata
Potosi
Cerrode Potosi
Potosi
Jakokkota
(=Jancocata)
Corocoro

leaves
leaves & fruits
fruit
fruits
leaves
leaves
leaves
leaves
leaves
leaves
pollen

Rondonia,Katira,Capoeira & RfoPreto

Serrados Carajas
Lagoa do Pires
Lagoa de Curuca
Morrode Itapeva,SP
Cata Altas
LagunaAngel& Laguna
Sardinas
Lake Pata-Hill of Six
Lakes
Morrodo Chapeu,Bahia
Umbuzerio
Lake Carajas & Lake
Salitre
Goias, Crominia
Bahia & Minas Gerais
Acre
Bahia
Minas Gerais,Fonseca
Minas Gerais,Fonseca
Bahia
Ouricanga,Alagoihas,
Bahia
Bahia
AlexandraFormation
near Curitiba,Parana
Pirabas,Para
SolimoesFormation
TaubateBasin
Sdo Paulo
TaubateBasin
Rio de Janeiro
Gandarela& Fonseca,
MinasGerais

pollen
pollen
pollen
pollen
pollen
pollen
pollen
pollen
pollen
pollen
leaves
leaves
pollen
pollen
leaves
leaves
leaves
leaves
leaves
leaves
leaves
fruit
pollen
leaves
pollen
leaves
pollen
leaves
leaves & fruit
pollen

Burnham& Graham
Historyof NeotropicalVegetation

Volume 86, Number2

1999

581

Appendix1. Continued.

Country

Geologicalage

Author(s)

De Lima,Junior&
Stefani
Brazil
Tertiary
Dolianiti
Brazil
Eocene-Oligocene Duarte& RezendeMartins
Brazil
Eocene-Oligocene Duarte& RezendeMartins
Brazil
Eocene
Fittipaldiet al.
Brazil
Cretaceous
Crane& Maisey
Brazil
Duarte
Cretaceous
Brazil
Aptian
Duarte& Silva-Santos
Brazil
Cretaceous
Herngreen
Brazil
Cretaceous
Osbornet al.
Brazil
Cretaceous
Pons et al.
Brazil& Guyana Quaternary
Van der Hammen&
Absy
Brazil,Ecuador, Tertiary
Langenheim& Beck
Colombia
Pleistocene
Colombia
Bakkeret al.
Colombia
Quaternary
Behlinget al.
Colombia
Quaternary
Gonzalezet al.
Colombia
Quaternary
Helmens& Kuhry
& Ran
Colombia
Quaternary
Hooghiemstra
Colombia
& Van
Quaternary
Hooghiemstra
der Hammen
Colombia
Van der Hammen&
Holocene
Gonzalez
Colombia
Van der Hammen&
Quaternary
Gonzalez
Colombia
Van der Hammen&
Quaternary
Gonzalez
Colombia
Pliocene-Present Andriessenet al.
Brazil

Colombia
Colombia
Colombia
Colombia
Colombia
Colombia

Eocene

Pliocene
Boureau& Salard
Pliocene-Present Hooghiemstra
Pliocene-Present Hooghiemstra
& Sarmiento
Pliocene
Howe
Pliocene
Sole de Porta

Date of
publication
1996

Local place name

Organsstudied

1955
1983

Macacu-Rifte
de Guana- pollen
bara
Recife
fruits
VargemGrandedo Sul
leaves & fruits

1985

VargemGrandedo Sul

leaves

1987
1991
1985
1993

Bacia de Sao Paulo

Santana
Santana
Maranhao

leaves
pollen
leaves
leaves

1975
1993
1992
1994

Alagoas
Santana(NE Brazil)
Pernambuco
Rondonia& Carajas

pollen
pollen
fruits& leaves
pollen

1968

Capanema& Para

ambers

1989
1999
1965
1986
1994
1993

PitalitoBasin
Pantanode Monica
SierraNevada del Cocuy
Paramode Agua Blanca

pollen
pollen
pollen
pollen
pollen
pollen

1960

1965

EasternCordillera:Lagu- pollen
na de La America
Sabana de Bogota:Pala- pollen
cio A
pollen
Boyaca

1993

EasternCordillera

1962
1994
1991

Bolfvar
EasternCordillera
Bogota

1974
1960
1997

leaves
MagdalenaValley
Cerrode la Popa, Carta- pollen
gena
pollen
High Plain of Bogota

1960

Cundinamarca

pollen& sediments
wood
pollen
pollen

1973

Sabana de Bogota

pollen

Colombia

Pliocene-Present Van der Hammen&

Hooghiemstra
Pliocene-Present Van der Hammenet
al.
Pliocene
Wijninga

1996a

CordilleraOriental

Colombia

Pliocene

Wijninga

1996b

High Plain

Colombia
Colombia
Colombia

Pliocene
Pliocene
Miocene

Wijninga& Kuhry
Wijninga& Kuhry
Berry

1990
1993
1936b

SubachoqueValley
Guasca Valley
Santander

pollen,leaves,
wood,seeds
pollen,seeds,
fruits,
wood,
leaves
pollen& leaves
pollen& leaves
leaves

Colombia

Annals of the
Missouri Botanical Garden

582

Appendix1. Continued.

Country

Geologicalage

Author(s)

Date of
publication

Organsstudied

Colombia

Mio-Pliocene

Colombia
Colombia

1965

TolimaProvince

Colombia
Colombia
Colombia
Colombia
Colombia

Hoorn
Miocene
Oligocenethrough Mirioni
Plioceneprobably
Mioceneor PlioPons
cene
Miocene
Pons
Miocene
Pons
"Late Tertiary"
Pons
Miocene
Pons
Sole de Porta
Miocene

1969
1976
1979
1985
1963

Colombia

Miocene

1996b

Colombia
Colombia

Oligocene-MioceneDuenas
late Eocene or Oli- Berry
gocene
Oligocene
Berry
Oligocene
De Porta& Sole de
Porta
Oligocene& Mio- Hoorn
cene
Oligocene
Marty
Oligo-Miocene
Pons
Sole de Porta
Oligo-Miocene

1977
1924a

wood
MagdalenaValley
Tolima
leaves
Hato Grande,Tolima
leaves
MagdalenaValley
leaves
Cira del Valle Formation,pollen
MagdalenaValley
Tequendama
pollen,leaves,
wood,fruits
pollen
PlanetaRica
Cundinamarca
fruits

1925b
1962

East of Bogota
MagdalenaValley

seeds
pollen

1988

Araracura

pollen

Colombia

Colombia
Colombia
Colombia
Colombia
Colombia
Colombia

Berry

Wijninga

1945b

Local place name

East of Cordillerade Bo- leaves

gotA
pollen
Caqueta River
Amaga,Prov.Bolfvar, wood
MagdalenaValley

1994a
1965

1933
near Yurumangui
& Naya leaves
1983
PlanetaRica
seed
1961a, b MagdalenaValley,San- pollen
tander
1924a
Cundinamarca& Bolivar fruits& seeds

Colombia
Colombia

Oligoceneand Eo- Berry

cene
Reid
Oligocene
Eocene, Miocene Berry

Colombia
Colombia
Colombia
Colombia
Colombia

Paleocene-Eocene
Eocene
Eocene
Eocene-Oligocene
Tertiary

Colombia
Colombia
Colombia
Colombia

Eocene-Oligocene Van der Hammen&

Garcia de Mutis
Van der Kaars
Paleocene
Paleogene
Doubinger& Pons
Cretaceous
Huertas

1983
1970
1960

Colombia
Colombia

Cretaceous
Cretaceous

Colombia

1933
1929c

1970
Doubinger& Pons
Gonzalez-Guzman 1967
Huertas
1977
Schuler& Doubinger 1970
1961b
Sole de Porta

Huertas
Huertas& Van der
Hammen
Colombia
Cretaceous
Stough
Colombia
CretaceousthroughVan der Hammen
lowerTertiary
Colombia& Guy- Quaternary
Wijmstra& Van der
ana
Hammen
Colombia,Brazil, Miocene
Hoorn
Peru

leaves

1966

& Naya fruits

near Yurumangui
Bolivar;Cundinamarca; leaves
Boyaca
leaves & fungi
MagdalenaValley
Tibu
pollen
nearMedellin
leaves
pollen
D'Amaga
near Monteria& Planeta pollen
Rica
pollen
pollen
cuticle
leaves

1969
1953

GuajiraProvince
Guajira
Sabana de Bogota,Zipaquira
Cundinamarca
Villeta

fruit
fruit

1968
1954

Sabana de Bogota
High Plain of Bogota

pollen
pollen

1966

LagunaAgua Sucia-Ru- pollen

pununi
Pebas, Solimoes
pollen

1994b

Volume 86, Number2

1999

Burnham& Graham
Historyof NeotropicalVegetation

583

Appendix1. Continued.

Country

Geologicalage

Author(s)

Date of
publication

Local place name

Organsstudied

Colombia
Costa Rica
Costa Rica
Costa Rica
Costa Rica
Ecuador
Ecuador

Miocene
Quaternary
Pliocene
Miocene
Miocene
Quaternary
Pleistocene

Hoorn& Lorente
Hooghiemstra
et al.
Graham& Dilcher
Berry
Graham
Bush & Colinvaux
Bush et al.

1992
1992
1998
1921a
1987
1988
1990

Caqueta River
Talamanca
Rio Banano nearZent
TalamancaValley
Uscari
Lake Ayauch
Mera,San JuanBosco

Ecuador

Pleistocene

Heine

1994

Mera

Ecuador
Ecuador
Ecuador
Ecuador
Ecuador
Ecuador
Ecuador
Ecuador
Ecuador
Ecuador
Ecuador
Ecuador
Ecuador
Ecuador &
Colombia
Guyana

Pleistocene
Miocene
Miocene
Miocene
Miocene
Miocene
Mio-Pliocene
Mio-Pliocene
Miocene
Mipcene
Eocene
Eocene
Cretaceous
Miocene

Liu & Colinvaux

Berry
Berry
Berry
Berry
Berry
Brown
Brown
Burnham
Wolf& vomRath
Berry
Berry
Shoemaker
Engelhardt

1985
1929a
1933
1934a
1935b
1945a
1946
1956
1955a, b
1876
1929d
1932
1982
1895

Mera
Loja
Loja Basin
Cuenca Basin
Malacatos
Loja Basin
Esmeraldas
PuntaGorda
Loja
Loja
Santa Elena
Santa Elena
El Oro
Loja & Santa Ana

1966

Panama
Panama

Quaternary
Quaternary

Bush & Colinvaux

Bush & Colinvaux

1990
1994

Panama

Pleistocene

Bush et al.

1992

Momnear Georgetown,
baka
El Valle
Darien-Lake Wodehouse
VeraguasProvince

Panama
Panama
Panama
Peru
Peru
Peru
Peru
Peru

Oligocene
Miocene
Eocene
Quaternary
Miocene
Pliocene
Pliocene?
Miocene

1918
1921b
1985
1994
1920a
1923
1925a
1985

Canal Zone
Canal Zone
Alcalde Dfaz
Andean
Tumbes
Cajamarca
Loreto
TumbesProvince

pollen,phytoliths
leaves & wood
nut
pollen
pollen
leaves
leaves
leaves
pollen

1992
1924b
1929b
1934b
1937e
1988
1995

Tumbes
ParinasPoint,Piura
Belen
ChiraValley
Piura
Morerilla
Laguna Baja

pollen
fruits
fruits& seeds
seeds
leaves
wood
pollen

1996
1925c
1937a
1937b

San Sebastian
Siparia& Moruga
ForestReserve,Fyzabad
Mud PlantForestReserve

pollen,leaves
leaves
leaves
leaves

Peru
Peru
Peru
Peru
Peru
Peru
Peru/Ecuador
PuertoRico
Trinidad
Trinidad
Trinidad

Paleocene-Eocene Leidelmeyer

Berry
Berry
Graham
Hansen et al.
Berry
Berry
Berry
&
Cruzado-Casteneda
Celi-Navarrete
Miocene
Woodet al.
Oligocene
Berry
Eocene
Berry
Oligocene
Berry
Eocene
Berry
Mourieret al.
Cretaceous
Pleistocene/Holo- Hansen & Rodbell
cene
Graham
Oligocene
Miocene
Berry
Miocene
Berry
Mio-Pliocene
Berry

pollen
pollen
pollen
leaves
pollen
pollen
pollen,phytoliths
pollen,radiocarbon
pollen,wood
leaves
fern
leaves
leaves
leaves
seeds
seeds
fruit
leaves
fruits
fruits
leaves & wood
leaves
pollen
pollen
pollen

Annals of the
MissouriBotanical Garden

584

Appendix1. Continued.

Country

Geologicalage

Author(s)

1985
1990
1987
1991

Venezuela
Venezuela
Venezuela
Venezuela

Quaternary
Holocene
Holocene
Holocene

Venezuela
Venezuela
Venezuela
Venezuela
Venezuela

1992
Holocene
Rull
Holocene
Rull
1996a
Pleistocene-RecentRull
1996b
1998b
Pleistocene
Rull
Holocene
Rull & Vegas-Vilarru1993
bia
Holocene
1987
Rull et al.
Pleistocene
Salgado-Labouriau 1987
Quaternary
Salgado-Labouriau 1991
Pleistocene
1988
Schubert& Rull
Miocene
1920b
Berry
Miocene
1921c
Berry
Miocene
1937c
Berry
Pliocene?
1939b
Berry
Miocene
Hambaleket al.
1994
1931
Miocene
Hoffman
Eocene and Mio- Berry
1936a
cene
Eocene
1939b
Berry
Eocene
Colmenares& Teran 1993
Eocene
De Di Giacomo&
1987
Van Erve
Eocene
Norem
1955
Eocene
Rull
1998a
Miocene
1924
Hollick

Venezuela
Venezuela
Venezuela
Venezuela
Venezuela
Venezuela
Venezuela
Venezuela
Venezuela
Venezuela
Venezuela
Venezuela
Venezuela
Venezuela
Venezuela
Venezuela
WestIndies

Leyden
Rinaldiet al.
Rull
Rull

Date of
publication

Local place name

Organsstudied

Lake Valencia
GranSabana
VenezuelanAndes
Guaiquinima,Chimanta
& Auyan
GranSabana
PantepuiProvince
Lake Valencia
Mesa de Caballo
Carinapay

pollen
pollen
pollen
pollen

PiedrasBlancas
PiedrasBlancas
MeridaAndes
Auyantepui
near Lake Maracaibo
Trujillo
Falcon
Anzoategui
Urumacotrough
Betijoque
Trujillo;Zulia; Falcon;

pollen
pollen
pollen
pollen
seed
leaves
leaves & fruit
leaves
pollen
leaves
leaves

MaracaiboBasin
MaracaiboBasin
Falcon

leaves
pollen
pollen

pollen
pollen
pollen
pollen
pollen

pollen
MaracaiboBasin
Misoa Formation
pollen
all islandsoftheWest leaves
Indies mentioned
separately

Volume86, Number2
1999

585

Burnham
& Graham
History
ofNeotropical
Vegetation

Appendix2. Comparisonof principalpalynomorphs

fromQuaternary
depositsin southernCentralAmericaand
1973; 2 = Bush &
northern
SouthAmerica.Numbersreferto the following
references:1 = Bartlett& Barghoorn,
Colinvaux,1988; 3 = Bush & Colinvaux,1990; 4 = Bush et al., 1990; 5 = Colinvaux,1996; 6 = Hooghiemstra,
1984 (table2); 7 = Hooghiemstra
et al., 1992; 8 = Hooghiemstra
& van der Hammen,1993; 9 = Horn,1985; 10 =
Liu & Colinvaux,1988; 11 = Martin,1961; 12 = van der Hammen& Absy,1994. Cheno-Am= ChenopodiaceaeTaxa are presented
Amaranthaceae
undifferentiated;
ERA = Euphorbiaceae-Rutaceae-Anacardiaceae
undifferentiated.
in thefollowing
order:fernsand fernallies, gymnosperms,
and dicots.
monocots,

Taxon
1
Acrostichum
Cnemidaria1, 7, 9
Ctenitis1
Cyatheaceae:Cyathea1, 3, 6, 7, 8, 9, 11
Danaea-type1
Elaphoglossum7
Hymenophyllaceae:
Hymenophyllum
7, 9
Isoetes7, 8, 11, 12
Jamesonia7
Lophosoria7
1, 7, 8, 11
Lycopodium
Monoletefernspore 1, 3, 4, 7, 8, 9, 10, 12
Ophioglossaceae7, 9
Pteris9
Triletefernspore 1, 4, 5, 8, 9, 10, 12
Selaginella 1
Araucaria5
Pinus 9
Podocarpus1, 4,5,6,7,8,9,10,11,12
Cyperaceae1, 2, 3, 4, 5, 7, 8, 9, 10, 11, 12
Eichhornia12
Eriocaulaceae7
Gramineae1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12
Iriartea1
9
Iriartea-type
Liliaceae 9
Mauritia5, 10, 12
Palmae 2, 4, 5, 7,9, 10, 12
Phytelephas1
8
Potomogeton
Sagittaria12
Typha1, 3, 7, 9
9
Abutilon-type
Acalypha1, 2, 6, 7, 8, 9, 10, 12
Alchornea1, 2, 4, 5, 6, 7, 8, 9, 10, 12
Alnus1, 4, 5, 6, 7, 8, 9, 11
AlfaroalOreomunnea
7, 9, 11
Alternanthera
3, 5, 9
10
Ambrosia-type
Anacardiaceae1, 8, 9
9
Anacardium-type
Annonaceae1
7
Antidaphne
Apeiba1, 9
Aphelandra1, 9
Apocynaceae1, 12

Lowland
Central
America
x
x
x
x
x
x

x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x

Upland
Central
America

Lowland
South
America

Upland
South
America

x
x

x
x
x
x
x
x
x
x
x
x

x
x
x
x
x

x
x
x
x
x

x
x
x

x
x

x
x

x
x
x
x
x
x
x
x
x

x
x

x
x
x

Annals of the
MissouriBotanical Garden

586

Appendix2.

Continued.

Taxon
Avicennia1, 9
Bauhinia emarginata1
Billia colombiana6, 8
Bocconia6, 8
Bombacaceae:Bombacopsis1, 12
Borreria1, 8
Bravaisia9
Brunellia6, 8
1
Byttneria
Bursera1, 5, 9
Byrsonima1, 3, 5, 9, 10
Calliandra 1
Canavalia 1
9
Cardiospermum
Caryophyllaceae
5, 7, 8, 9
Cassipourea1
Catopsis1
Cavanillesia1
Cecropia3, 5, 6, 7, 8, 9, 10, 12
Celtis2, 9, 10, 12
6
Cestrum
Cheno-Am1, 3, 8, 9, 10
Cissus1
Clethra6
Clusia C.-type7, 8
1
Coccoloba-type
Combretaceae-Melastomataceae
3, 9
1
Combretum
Compositae1, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12
1
Compsoneura
Conocarpus1
Copaifera1, 5
Cordia6, 9, C. alliodora1
Coriaria8
Couroupita1
Croton:Crotonoideae1, 7, 8, 9
Cruciferae7, 8
Cupania 1
Cuphea 12
Curatella12
Daphnopsis6, 7
Dialyanthera1
Didymopanax2, 7, 10, 12
Dodonaea 6, 7, 8
Dorstenia9
1
Drepanocarpus
lunatus-type
Drimys6, 7, 8, 9, 11
ERA 9, 10, 12
Ericaceae 1, 4, 5, 6, 7, 8, 9, 11
1
Erythrina
Escallonia 7
Eugenia 9

Lowland
Central
America

Upland
Central
America

Lowland
South
America

x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x

x
x
x

x
x
x
x
x
x
x
x

x
x
x
x

x
x

x
x

x
x
x
x
x
x
x
x
x
x

Upland
South
America

x
x

x
x

x
x

x
x
x
x
x
x

x
x

x
x
x
x

x
x
x

x
x

x
x
x
x

Burnham& Graham
Historyof NeotropicalVegetation

Volume 86, Number2

1999

Appendix2.

587

Continued.

Taxon
Euphorbia8
Ficus 2, 9, 10, 12
Faramea 1
Fuchsia 7, 8
Gaiadendron6, 7, 8
Galium8
Gentianaceae7
Geranium7, 8
1
cf. Glycydendrum
G. amazonicum1
9
Gomphrena
Guazuma9
Gunnera7
1
Gryanthera
darienensis-type
Hedyosmum
1, 3, 4, 5, 6, 7, 8, 9, 12
Heliocarpus6, 7, 8
Hesperomeles
6, 7
Hibiscus9
Hieronyma1, 6, 7, 8, 10
Hippocrateavolubilis1
Humiria5
Hura 3, 5
Hydrangea1
8
Hydrocotyle
Hydrophila9
Hypericum
6, 7, 8
Ilex 1, 4, 5, 6, 7, 8, 9, 12
Ipomoea1
Juglans6, 8, 9
Jussiaea1
Justicia1
Labiateae8, 12
Lafoensia1
Laguncularia9
5
Lecythidaceae-Eschweilera
Leguminosae2, 4, 5, 8, 10
Caesalpinioideae5
Mimosoideae1, 9, 10
1
Leucaena multicapitula
Licania arborea1
Limnocharis
3
Liquidambar9
Liriosma1
Loranthaceae7, 12
Luehea 1, 9
Lysipomia7
Lythraceae8
Mabea 1
5
Machaerium-Dalbergia
Malouetia1
Malpighiaceae1, 8, 9, 10, 12

Lowland
Central
America
x
x

x
x
x
x
x
x
x
x
x

Upland
Central
America

x
x
x

x
x
x
x
x

x
x
x
x

x
x
x
x

x
x
x

x
x
x
x
x

x
x
x

x
x

x
x
x
x
x
x

Upland
South
America

x
x
x

x
x

x
x
x

Lowland
South
America

x
x
x
x

x
x
x

Annals of the
MissouriBotanical Garden

588

Appendix2.

Continued.

Taxon
Malvaceae 1, 8, 9
Manihot1
Maripa 1
Melastomataceae-Combretaceae
1, 2, 4, 5, 6, 7, 8, 10
Meliaceae-Sapotaceae1, 3, 5, 9, 12
Mendoncia1
Menispermaceae1
Merremia1
Miconia6, 8, 12
Montia7
Moraceae,Urticaceae,Urticales1, 2, 3, 4, 5, 6, 7, 8, 9, 10
Mortoniodendron
9
Mucuna 1
Mollugo1
Myrica1, 3, 5, 6, 7, 8
8
Myriophyllum
Myrrhiodendron-Niphogenton
7
Myrtaceae1, 3, 4, 5, 6, 7, 12
Ochroma1
Oreopanax7
1
Oryctanthus
Ouratea0. guatemalensis1, 12
Onagraceae9
Pachira 1
Pacouria 1
Paullinia 1
Panopsis8
Pelliceria9
Phyllanthus1
Pilea P trema2, 5, 6, 8
Piper9, 10
Piperaceae 1, 2, 4, 5, 7
Plantago 7, 8, 9
Polygalaceae1, 6, 8, 12
Polygonum1, 7, 8, 9
Polylepis-Acaena
6, 8
Pontederiaceae1
Portulacaceae1, 8
Proteaceae4, 5, 6
Pseudobombax1
Psidium9
1
Pterocarpus-type
Puya 7
Quararibea1
Quassia 1
Quercus3, 5, 6, 7, 8, 9, 11
Randia 1
Ranunculaceae7, 8, 12
Rapanea R.-type5, 6, 7, 8, 9
Rauwolfia1
Relbunium8
Rhizophora1, 9

Lowland
Central
America
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x

Upland
Central
America

x
x
x
x
x
x
x

x
x

x
x

x
x

x
x

x
x

x
x
x

x
x
x
x

x
x
x
x
x

x
x
x
x
x

Upland
South
America
x

x
x
x
x
x
x
x

Lowland
South
America

x
x

x
x

x
x

x
x

Volume 86, Number2

1999

Appendix2.

Burnham& Graham
Historyof NeotropicalVegetation

589

Continued.

Taxon
Rosaceae 9, 10
Rubiaceae 1, 7, 12
Rumexcostaricensis
7
Salix-S.-type7, 9
Sapium 1, 2, 7, 8, 12
S.-type1, 9
Sapotaceae 5
8
Scrophulariaceae
Sechiumedule-type1
Solanaceae 7, 8, 10, 12
Spondias-type
9
Styloceras6, 8
1
Swartziapanamensis-type
Symphonia1, 4, 5, 9, 12
SymplocosSymplocaceae1, 5, 6, 8
1
Tabernaemontana
Tetracera1
Thalictrum
T-type8, 9
Tiliaceae 4, 5
9
Tournefortia
Trema2,5,9,10
1
Trichanthera
Tristicha1
Ulmus7, 9
Umbelliferae
9, 10, 11
Utricularia1
Valeriana7
Vallea 6
Viburnum
6, 7, 8
Virola1, 9
Vochysia1
Weinmannia
W.-type
4, 5, 6, 7, 8, 9, 10, 12
Zanthoxylum
1, 9
Zea 1,2,10

Lowland
Central
America
x
x
x
x
x
x
x

Upland
Central
America
x
x
x
x

x
x
x
x
x

Upland
South
America

x
x
x
x
x

x
x
x
x

x
x
x
x
x
x
x
x
x
x
x
x
x

Lowland
South
America

x
x

x
x

x
x
x

x
x

x
x