Neotropical Floristic Diversity A, Gentry

Neotropical Floristic Diversity: Phytogeographical Connections Between Central and South
America, Pleistocene Climatic Fluctuations, or an Accident of the Andean Orogeny?

Author(s): Alwyn H. Gentry
Source: Annals of the Missouri Botanical Garden, Vol. 69, No. 3 (1982), pp. 557-593
Published by: Missouri Botanical Garden Press
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NEOTROPICAL FLORISTIC" DIVERSITY:
PHYTOGEOGRAPHICAL CONNECTIONS BETWEEN
CENTRAL AND SOUTH AMERICA, PLEISTOCENE
CLIMATIC FLUCTUATIONS, OR AN ACCIDENT
OF THE ANDEAN OROGENY? ,2
ALWYN H. GENTRY3
The interchangeofplantspeciesbetweenNorthand SouthAmericahas been

a majorfactorin determining the Neotropicalphytogeographicalpatternsob-
servedtoday.Although thishas longbeen realized,recenthistoricalgeological
evidenceas wellas increasingknowledgeof today'sdistributionalpatternsnow
makespossiblea morethorough analysisofhow thesepatternsoriginated. This
paperbrieflyreviewsthegeologicalbackground, summarizes thecomposition of
theextantNeotropicalflora,pointsoutthestriking
ecologicalconsistencyofmany
taxa and lifeforms,and suggestshow some of the presentphytogeographical
patternshave developedfromtheinterplay ofthesefactors.
GEOLOGICAL BACKGROUND
Severalmajorgeologicaleventshave had profoundeffectson the evolution

and distributionoftheLatinAmericanflora.One ofthemostsignificant ofthese
was the separationof SouthAmericafromAfricathatbegan only 127 million
yearsBP (Rabinowitz,1976;McKenna,1981)withcontactor near-contact pres-
entuntil80-90millionyearsBP (Raven& Axelrod,1974;McKenna,1981).Dur-
ingmostofthefirstthirdofangiosperm evolution,a timeduringwhichmanyof
themodernordersandfamiliesofplantsarose,SouthAmericawas a partofthe
WestGondwanaland cradleoftheangiosperms (Raven& Axelrod,1974).How-
ever,duringvirtuallyall oftheTertiaryandmuchoftheCretaceousSouthAmer-
ica was an islandcontinent,at leastfromtheperspectiveoftropicalplants,and
mostoftheevolutionofitsrichand variedfloratookplace in isolationfollowing
separationfromAfrica.Thus,suchcharacteristic andecologicallyimportant
trop-
1 Thispaperwas presented inbothSymposiapublished inthisissueoftheAnnalsoftheMissouri

BotanicalGarden-theSymposium on PlantGeographical Resultsof ChangingCenozoicBarriersat
theXIII InternationalBotanicalCongressin Sydney,Australia,1981;andthe1981Systematics Sym-
posiumof theMissouriBotanicalGarden.As used herein,references documented as "thissympo-
sium"includepapersfrombothsymposia.
2 Thispaperstemslargely frominsightsdevelopedduringfieldworkovera numberofyearsin
variouspartsofLatinAmerica,and supported by theNationalScienceFoundation (GB 40103,INT-
7920783,DEB-8006253, DEB-UT-20325, DEB-8006253), theNationalGeographicSociety,andUSAID
(DAN-5542-G-SS-1086-00). I thankP. Ashton,D. Axelrod,C. Dodson,A. Graham,P. Raven,J.
Rzedowski,J. Terborgh, and B. Simpsonforreviewing the manuscript and numerouscolleagues
includingD. Austin,H. Balslev,C. Berg,W. Burger, L. Constance,G. Davidse,R. Faden,P. Fryxell,
S. Graham,R. Haynes,M. Huft,N. Holmgren, M. Johnston,J.Kuijt,J.Luteyn,M. McKenna,W.
Meijer,J. O'Neill, R. Bleiweiss,T. Plowman,M. Poston,J. Pringle,H. Sleumer,L. B. Smith,D.
Soejarto,C. Stace,P. Taylor,W. Wagner,D. Wasshausen, H. Wilson,andD. Wunderlin forprovid-
ingdataon Neotropical speciesnumbers or distribution
patterns
oftheirtaxonomic specialties.
3 MissouriBotanicalGarden,P.O. Box 299,St. Louis, Missouri63166.
ANN. MISSOURI BOT. GARD. 69: 557-593. 1982.
.75/0
0026-6493/82/0557-0593/$03

558 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 69
ical Americanangiosperm familiesas Bromeliaceae,Humiriaceae,Cactaceae,

and Caryocaraceae,had ampletimeto evolveand radiatein SouthAmerica.
Although thehistory oftheseparationofSouthAmericafromAfricais clear,
the tectonicsof the earlierseparationof NorthAmericafromthe Gondwanan
landmassis stilluncertain (see Coney,thissymposium). Apparently NorthAmer-
ica (includingthe CentralAmericanpeninsula)separatedfromAfrica-South
Americain Jurassictime,wellbeforetheoriginoftheangiosperms (Lillegraven
et al., 1979).However,by late Cretaceousa chainof volcanicislands(Nicoya
Complex)haddevelopedconnecting CentralAmericawithEcuador(Dengo,1975;
Lillegravenet al., 1979).New geologicalevidence(Dickinson& Coney, 1980)
suggeststhata connection betweennuclearCentralAmericaand SouthAmerica
was reestablished inLate Cretaceousas bothmajorAmerican landmassesmoved
westwardmoreor less in tandem.
How muchof the late CretaceousconnectionbetweenCentraland South
Americawas above sea levelis notknown,although itprobablyconsistedmostly
of an interrupted islandarc. Furthercomplicating the picture,muchof north-
westernSouthAmericawas submerged duringmostof the Cretaceous(Irving,
1975).Boththelow coastalrangeofwesternColombiaand southern Darienand
thenorthern AndeanCordilleraOccidentalwereoriginally islandarcs associated
withwestwardmovement of the SouthAmericanplate duringupperMesozoic
time(Zeil, 1979: 193). McKenna (1981) has suggestedthatthe presentPacific
coast of northern SouthAmericamayhave resultedfromaccretionduringthe
earlyTertiary ofleft-behind fragments ofthesouthend oftheCentralAmerican
volcanicarc. Accordingto McKenna(1981) some oftheislandsofthisarc may
have remainedabove water,separatedfromNorthAmericaonlyby sequential
openingand closingof watergaps,untilcollidingwithSouthAmericaand pro-
vidinga plausiblescenariofor"Noah's Arc" dispersalofhystricognath rodents
betweenNorthand SouthAmerica,a modelconsistentwiththe discoveryby
Juteauet al. (1977) thatmuchof the west coast of Ecuador is formedfroma
blockof originally oceaniccrust.
Therewas a generalregression ofepicontinental
seas fromnorthwestern South
Americaat theendoftheCretaceous(Harrington, 1962;Lillegraven et al., 1979)
concomitant witha late Cretaceousorogenythatgave riseto theforerunner of
theColombianCordillera Occidental(Irving,1975).Morethan160kmwas added
to thenorthwestern SouthAmericancontinental marginduringLate Cretaceous
time.The GreaterAntilles,thenfarto thesouthoftheirpresentpositionand in
partsubmerged, presumably also constituted
partofsomekindoflateCretaceous
inter-American connectionas suggestedby'Malfaitand Dinkelman(1972) and
discussedin the contextof biogeography by Tedford(1974) and Rosen (1974).
(See Pregill(1981)fora different butmuchofthegeologyon which
interpretation
it is based (Perfit& Heezen, 1978)has apparently been superseded(Dickinson
& Coney, 1980;Coney,thissymposium).) Accordingto thisinterpretation, the
continued westwardmovement ofSouthAmericaandtheslightly southwestward
movementof NorthAmericaled to decouplingfaultsthatseparatedtheproto-
Antillesfrombothland masses. A new subductionzone thenformedto their
northeast as theproto-Antilles movednortheastward. Continued westwardmove-
mentof SouthAmericaand southwestward movement ofNorthAmericaled to

1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 559
pinchingoffof a segmentof thePacificplateas the Caribbeanplate,withrela-

tivelyeastwarddisplacement of the whole Caribbeanregionrelativeto South
America.ProbablynuclearCentralAmericasouthoftheMotaguafaultalso moved
eastwardwiththeCaribbeanplatefroman originalpositionmoredirectlysouth
ofMexico,closingpartofthegap betweenNorthand SouthAmerica(Dickinson
& Coney,1980).In thelateTeritary, formation oftheCentralAmerican trenchand
an associatednewepochofvolcanismagainled to upliftofislandsin theregion
betweenSouthAmericaand nuclearCentralAmerica.These islandseventually
coalesced intotoday'slowerCentralAmericawithsubstantial land connection
acrosstheIsthmusofPanamaestablishedin thePlioceneapparently onlyabout
3 millionyearsago (Keigwin,1978;Marshallet al., 1982).
Although closingoftheIsthmusofPanamabetweenNorthandSouthAmerica
was clearlyone ofthemostimportant "changingCenozoicbarriers"fortheLatin
Americanbiota,thepresenceof a previous,albeitinterrupted, late Cretaceous
connection wouldhaveprovidedshorter watergapsbetweenthecontinents fairly
earlyand thusmaymodifyour conceptofthephytogeographical importance of
thislateevent.In lateCretaceoustimemanymodernfamilies andgeneraofplants
wereextantin WestGondwanaland and in a positionto takeadvantageofisland
stepping-stones betweenSouthand NorthAmerica.
Thus mostof the cases of exclusivelyor predominantly Americanfamilies
likeCactaceaewithstrongly differentiatedcomponents inbothtropicalNorthand
SouthAmerica,whichRaven and Axelrod(1974) accountedforby chancerela-
tivelylong distancedispersalat varioustimesduringthe earlyCenozoic,may
reflectinsteadmoreor less directlate Cretaceousmigration betweenthe two
continents, presumably mostlyvia islandhopping.Similarly, the dichotomous
compositionof the West Indianflora,withboth strongsouthernand northern
affinities,would be expectedas the resultof an originalstockingof a proto-
AntilleanregionlocatedbetweenNorthand SouthAmericaas emphasizedby
Rosen (1974),althoughhis arguments fordirectland connections do notaccord
withthegeologicalevidence(Coney,thissymposium). Thisdichotomy mayhave
beenevenstronger to
intheearlyTertiary judge from thepresence ofnow locally
extinctotherwise exclusivelySouth American taxa likeAetanthus (although there
may be identificationproblems in separating thisfrom thegeneralized northern
Loranthuspollentypethatwas alreadypresentin Europe in the Eocene, ac-
cordingto Muller,1981)and Catostemmain theOligoceneofPuertoRico (Gra-
ham & Jarzen,1969)alongwithnorthern elementslikeFagus, Hauya, Engel-
hardtia,Liquidambar,and Nyssa, whichhave neverbeen able to reachSouth
America.
Thattheinterrupted linkbetweenNorthand SouthAmericawas completely
ruptured at thebeginning oftheTertiary, as theproto-Antilles begantheirnorth-
wardmovement, meansthatonlythoseveryold taxa alreadyextantin theCre-
taceouswouldhave had the opportunity formoreor less directinter-American
islandhoppingpriorto the Plioceneformation of the Isthmusof Panama. In
general,thefloraoftheearlyTertiaryofwhatis now temperate NorthAmerica
seemsto havebeenmorelikethemoderntropicalAmericanflorathanitis today.
However,it is verydifficult to judge to whatextentthe apparentaffinities of
Paleoceneand earlyEocene NorthAmericanand SouthAmericanflorasmight

representshareddescentfromhypothetical commonwide-ranging middleCre-

taceousearlyangiosperm stocks,or whethertheysuggestan independent, rela-
tivelydirectfloristicinterchange betweenSouth Americaand tropicalNorth
Americasubsequenttotheseparation ofAfricaand SouthAmerica.The difficulty
offloristic
interpretationis compounded by theserioustaxonomicerrorsin iden-
of fossilflorasthatare now knownto have characterized
tification muchearly
paleobotanicalwork(cf.Graham,1972:8; Dilcher,1974;Hickey& Wolfe,1975).
As summarized by Leopold and MacGinitie(1972),thefloristic of the
affinities
RockyMountainregion,theonlypartofthenbioticallytripartite NorthAmerica
in potentiallydirectcontactwithnuclearCentralAmerica,show successive
changesthroughthe earlyTertiary.They findthe generalizedPaleocene flora
to relateto thatof any extantphytogeographic
difficult region,but suggestthat
themesic earlyEocene florawas primarily relatedto the southeastAsian sub-
tropicaland warm-temperate mixedmesophytic forest,themiddleandlateEocene
florasto thoseofthesubhumid CentralAmericantropics,theOligoceneflorato
the live oak forestsof the highlandsof northern Mexico, and a finalshiftto
modernCordilleran conifer forestsintheMiocene(Leopold& MacGinitie,1972).
Presumablymost of the generasharedwithSouth Americaduringthe latest
Cretaceousand earliestTertiary reflectwide-rangingCretaceouscommonances-
tors(cf. Raven & Axelrod,1974),whilethe subhumid middleand late Eocene
floraanalyzedas characteristically and autochthonously tropicalAmericanin
originwouldhavetorepresent eitheran olduniquelytropicalNorthAmerican flora
(which later must have spread to South America to account fortheoverwhelm-
inglypan-American distributions ofmostofthosegeneratoday)or earlyfloristic
interchange withSouthAmerica.Contrary totheCentralAmerican4 distributional
centers suggestedfor such genera as Ocotea, Beilschmiedia, Cedrela, Luehea,
Oreopanax,and Swartziaby Leopold and MacGinitie(1972),all are betterrep-
resentedtodayin SouthAmericathanin CentralAmericawiththepossibleex-
ceptionof Cedrela(threeof theeightspeciesonlyin CentralAmerica,and two
othersalso occurringthere).Even threeof the fourgenera-Homalium,Ber-
noullia,Beilschmiedia,and Engelhardtia-related to Rocky Mountainfossilforms
and suggestedas endemicto CentralAmericain theirAmericanranges,are as
well or betterrepresented in SouthAmerica.Unfortunately theTertiaryfossil
recordfromCentralAmericaand SouthAmericais inadequateto show when
suchgeneraachievedtheirmodernpan-neotropical butphytogeo-
distributions,
graphicand ecologicalevidencesuggeststhatit was longbeforeestablishment of
thePanamanianlandbridge.Manyoftheseplantshave wind-dispersed or mam-
mal-dispersed seeds or fruitsand are unlikelycandidatesforlongdistancedis-
persal(Gentry,1982a),whichwouldbe consistentwithhavingachievedtheir
presentdistributions directinter-American
via a relatively route,pre-
migration
sumablypriorto theEocene.
Althoughsome SouthAmericanfloristic elementsevidently reachedCentral
Americaearly,perhapsvia islandhoppingalongthe late CretaceousAntillean
connection,the-Tertiary florasof SouthAmericaand NorthAmericaremained
thispaperin a broadsenseto includeMexico.

4Central Americais usedthroughout

fundamentally different
(Germeraadet al., 1968).For exampleGraham's(1973,
1982) analysisof the Paraje Solo formationsuggeststhatin the Miocene the
southern Veracruzlowlandshad onlya fewscattered representativesoftheSouth
American-derived tropicalforestwhichnow characterizesthe region,whilea
basicallyNorthAmericanplantcommunity, similarto thedeciduousforestofthe
easternUnitedStatesand todayrestricted to intermediateelevations,was well
represented.Pollenof ten of the fourteenarborescent generathatwouldhave
been expectedin similardepositsin the southeastern UnitedStatesis present.
Whileredeposition ofpollenfromplantsgrowingat higheraltitudescouldhave
affectedtheseresults(Axelrod,pers. comm.),it is clearthattheeasterndecid-
uous forestelementswereat least presentin theregion.In contrast,the Pale-
ocene pollenfloraof Colombiacontainedexclusivelysuchlowlandtropicalele-
mentsas Annonaceae,Bombacaceae,Melastomataceae,Araceae, and several
generaofpalms:Mauritia,Astrocaryum, and severalpollengenera(Hammen&
Garcia,1966).
Furthersouthin CentralAmerica,in thenrecentlyemergedPanama,only
three of the temperateNorth American genera-Alnus, Juglans, and Myrica-
are presentin the Miocene Gatunformation and none of themare presentin
earlierPanamaniandeposits(Graham,1973).These samethreegeneraappearin
theSouthAmericanpalynological recordonlysubsequentto closingoftheIsth-
mianconnectionin latestPlioceneand Pleistocenetimes.The manywidespread
SouthAmericanfamiliesand generathattodaybarelyenterCentralAmericain
easternPanamaprovideevidencethattheTertiary barrierto northwardmigration
was equallyeffective. As mightbe expected,none of theSouthAmericantaxa
thatbarelyenterCentralAmericaare represented in theWestIndieseither.To
summarize, thewatergapbetweenNorthand SouthAmericaseparatedtwovery
distinctCenozoicflorasinthetwocontinents despitea probableearlyopportunity
forrelativelydirectislandhoppingacross theproto-Antillean chain.However,
theopportunity forlimitedlate Cretaceousmigration betweenSouthand North
Americawouldreadilyaccountforthemanycharacteristically Gondwanantaxa
withdistinctive and stronglydifferentiated
CentralAmericanderivatives(see
below).
The secondCenozoicgeologicaleventwithmajorphytogeographical impor-
tancefortheNeotropicswas theupliftoftheAndes.Although theAndeanorog-
enywas certainly morecomplicated thanoftensupposed(Zeil, 1979),thegeneral
pictureof majorupliftof the alreadyextantsouthernand centralAndes in the
Mid-Cenozoicand ofthenorthern Andesmorerecentlyseemswell established.
The ColombianCordilleraCentralis olderand existedalreadyin theCretaceous
(Zeil, 1979: 109)butprobablywas erodeddownto a low rangeof hillspriorto
therecentorogeny.Most of theupliftofthenorthern Andestookplace onlyin
thelastfivemillionor so years,duringPlioceneand Pleistocenetimes(Hammen,
1974;Flenley,1979).The Andesare unique:by farthemostextensivemountain
rangein theworld'stropics.
A thirdeventof majorphytogeographic importance was theadventofPleis-
toceneclimaticfluctuationsassociatedwithglacialadvancesandretreatsat higher
latitudes.In themontanetropicsthese-climatic oscillationstooktheformof an
altitudinalloweringand compressionof vegetationalzones, as elegantlydocu-

562 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL.69
mentedby van der Hammen(1974) and his associatesfromthe palynological

recordin theColombianCordilleraOriental.In thelowlandtropicstemperature
changesassociatedwithglacialadvanceswere minimalbut changesin precipi-
tationwerepronounced. The cyclesofglacialadvanceandretreat wereassociated
in thetropicallowlandswithalternating dryand wetperiodsrespectively. Con-
traryto earlysuggestions (e.g., Haffer,1970),thewetperiodsofthetropicswere
notthesameas thewellknownpluvialperiodsofaridtemperate and subtropical
desertsthataccompaniedglacial advances.Instead,tropicalwet periodswere
perhaps180?out of phase withthe temperate-subtropical pluvialperiods;as if
the glacialadvancestiedup so much of the water of the earth's normalatmo-
sphericcirculation not
that enough was leftfor "normal" tropicalrainfall.As a
resultof these cyclicalchanges in precipitationthe lowland neotropical forests
wereperiodically reducedin extentto scatteredpockets,chiefly aroundthepe-
ripheryofAmazoniaduring dryperiods(Haffer, 1969,1978;Simpson,1971;Prance,
1973,thissymposium; Simpson& Haffer,1978).
The biologicalsignificance of thisdynamicmodel,verydifferent fromthe
popularconceptionofthe stable"forestprimaeval," is thatit providesoptimal
conditionsfor speciation,as populationsof tropicalforestspecies repeatedly
fragment and recoalesce(see papersin Prance,1982).Some problemswiththe
Pleistocenerefugemodelhave been noted.For example,have long-livedtrees
and lianashad adequatetimeto speciateso profusely in therelatively shorttime
and fewgenerations availablesincethebeginning of thePleistocene?Moreover
thedistributionalpatterns thathave been citedas evidenceforformer refugia-
speciespairsinmanyunrelated taxacomingtogether at coincident contactzones-
could also resultfromin situspeciationalongenvironmental gradients.Never-
thelessthePleistocenerefugemodelhas beenwidelyacceptedbybiogeographers
as a generalexplanationaccounting formuchtropicalspeciation.Not onlydid
forestspecies successfully survivedryperiodsin thesehabitatislands,butthe
repeatedcyclesofmultiple rangefragmentation couldpotentially have multiplied
theirnumbers.In otherwords,theNeotropicshave manyspeciesbecause ofthe
multiplicative effectof its more numerousrefugia;Africahas few because it
generally lackedsuchrefugia.
FLORISTIC BACKGROUND
One oftheoutstanding featuresoftheNeotropicalflorais itsextremerichness

in species.ThusRaven's(1976)estimatefornumberofNeotropicalplantspecies
was 90,000,threetimesas highas hisestimatefortropicalAfricaplusMadagascar
and 21/2timestheestimatefortropicalAustralasia.Prance(1977) and otherau-
thorshave generally acceptedRaven's estimates.Otherrecent,perhapsslightly
higher,estimatesforcontinental floristic are availableforthePalaeo-
diversity
tropics-30,000speciesforcontinental tropicalAfricaalone(Brennan,1979)and
25,000-30,000 speciesfortheFloraMalesianaregion(Jacobs,1974).Nevertheless
it seems clearthattheestimatednumberof plantspeciesfortheNeotropicsis
muchhigherthanfortheentirePalaeotropicalregion.Prance(pers. comm.)es-
timates30,000speciesforAmazonianBrazilaloneandthebestavailableestimate
forCentralAmericais 18,000to 20,000(Gentry,1978)and an (overlapping) one
forMexico is 14,000to 20,000(Rzedowski,1965,1978).

Whileit is generallyappreciatedthatAfricais floristically depau-

relatively
perate(Richards,1973;Brennan,1979),manystudying the floraof Southeast
Asia (Whitmore, 1975;Ashton,1964,1977)have suggestedthatthedipterocarp
forestsof thatregionare the world'srichest,citingsamplesof numberof tree
speciesover10cmdbhas evidence.Actually, Pranceetal.'s (1976)recentsample
-
of 179treespecies 15 cm dbh in a hectare of forestnearManausis as diverse
as mostoftheSoutheast Asian forests. If individualSoutheastAsianforestshave
as manyor moretreespecies as their neotropical equivalents, can theNeotropics
reallyhave as many more species as suggested by Raven's estimates?
The availableestimations ofNeotropicalfloristic diversityare verytentative.
In orderto assess moreaccurately whether theNeotropicsarereallyas incredibly
species richas suggestedby Raven's estimates,I decidedto tryto countthe
numberof speciesdirectly.First,I compileda listof all Neotropicalseed plant
generabased on the availableregionalflorasand familialmonographs, supple-
mentedby a searchthrough theentireMissouriBotanicalGardenherbarium for
additional genericnamesfromtheregion.Although a fewsmalllocal generawere
probablymissed,theresulting listof over4,200Neotropicalgeneraseemsrea-
sonablycomplete.Second,I estimated thenumberofNeotropicalspeciesineach
genus,usingrecentmonographs suchas theFloraNeotropicaseries,whentreat-
mentswereavailable,andthefigures suppliedby Willis'sDictionary (AiryShaw,
1973)inmostothercases. A fewadditional datasourceson speciesnumbers were
used forsomegroups-CompositaeweretakenfromHeywoodet al. (1977),Le-
guminosaemostlyfromPolhilland Raven (1981), OrchidaceaefromDressler
(1981), Solanaceae fromD'Arcy (1979), Verbenaceaeand Eriocaulaceaefrom
Moldenke(1980),Gramineae(mostlyDavidse,pers.comm.),and Bignoniaceae,
Sabiaceae, and Buxaceaefrommyowndata. The Willisfigures forspeciesnum-
bers are directlyusefulonlyforgenerarestricted to theNeotropicsand in the
fewcases wherespeciesnumbersfora genusare givenby geographical region.
The resultant data set accountedforover3,660generawithalmost65,000Neo-
tropicalspecies.
Unfortunately, theother533non-monographed non-endemic Neotropicalgen-
eraincludea totalof60,000speciesbased on theWillisfigures. Thesegenerafall
intotwomaingroups-largepantropical generalikePiperorEugenia,orbasically
LaurasianherbsthatrangesouthintothemontaneNeotropics.About20,000of
thesespeciesfallintopantropical generathattendto be wellrepresented in the
Neotropicssuggesting thata thirdto a half(i.e., 7,000-10,000)ofthemare Neo-
tropical.The remaining 40,000+ speciesare eitherin cosmopolitan genera(e.g.,
Ipomoea) or arebasicallytemperate zone generawithrelatively poorNeotropical
representation(e.g., Astragalus, Draba, Carex, Gentiana). Perhaps only 10-15%
ofthetotalspeciesof thesegeneraare Neotropical,implying an overalltotalof
at least76,000Neotropicalspecies.
However,thereis anotherseriousproblemincompiling suchan estimate.The
Neotropicsare generallymuch more poorly known thanotherparts
floristically
of the worldand verymanyNeotropical plant species remain undescribed.In
regionslikethewestern Colombian Choco, probablya quarter ofall plantspecies
endemic
are strictly and in many groupslike Araceae or Ericaceae almostnone
oftheendemicspecies have been described(Gentry,1982a). A well-documented

examplecomesfromtheRio Palenquefieldstationin westernEcuadorwhere,

in theprocessofwriting a local florula(Dodson & Gentry,1978),61 newspecies
were discoveredin a minuscule1.7 km2area; subsequentcollectionsand re-
evaluationof tentatively identifiedproblemtaxa now bringsthetotalto almost
100 species describedfrom Rio Palenque. Such figuressuggestthattheremust
be at least 10,000undescribedneotropicalplantspecies thatwouldelevatethe
totalnumberofneotropical seed plantspeciesto 86,000or more.
We mayconcludethatRaven's originalestimateof90,000neotropical species
was fairlyaccurate.Thattheneotropicsas a wholeare extraordinarily richerin
plantspeciesthanotherpartsoftheworldseemsestablished beyondanyserious
doubt.
The Neotropicalfloragenerallyshowsverystrongpantropicalconnections.
Thatpantropical distributions predominate amongtropicalfamiliesis wellknown
(e.g., Good, 1974).However,thesephytogeographic areevenstronger
similarities
thangenerally realized,especiallyat thegenericleveland especiallywithAfrica.
For example,30% of thegenerathatoccurat Makokou,Gabon,are also found
in theNeotropicsand manymoreAfricangeneraare morelikesomeNeotropical
genusthanlike otherAfricanones (Hladik & Halle, 1973; Gentry,in prep.).
Statisticalcomparisonsof percentagesof generain commonbetweendifferent
tropicalareas (e.g., Thorne,1973)are especiallymisleading sincetheimportant
generatendto be sharedwhilesmallsegregategeneracontribute disproportion-
atelyto thedifferences. The pictureis also confusedby taxonomicparochialism
on thepartofworkerson different continents.
The relevantpointof thesepantropical floristic in thecontextof
similarities
thisanalysisis thatthepeculiarities thatdistinguishthepresentNeotropicalflora
have arisendespitea commonfloristic stocksharedat least withAfrica(i.e.,
West Gondwanaland)and to a largeextentwithtropicalAsia as well. In this
contextit is almostirrelevant whetherthisintercontinental commonality was
derivedfromdispersaleventsor continental movements.The questionto be
addressedhereis whythissharedfloristic stockhas givenriseto so manymore
speciesin theNeotropics.
ENDEMIC FAMILIES
The greaterdiversityin the Neotropicsresultsin partfroma conspicuously

largerand moreecologicallyimportant complement of endemicfamilies.Even
broadfamilialdelimitations,
traditional
retaining thereare at least38 endemicor
endemic(i.e., withone or twoAfricanspecies,mostofwhoseances-
essentially
torsprobablyarrivedrelatively via longdistancedispersal)Neotropical
recently
familieswitha totalof 5,690species (Table 1). Thirteenof thesefamilieshave
over50 speciesas comparedto onlythreeendemicpalaeotropical familieswith
morethan50 species-Dipterocarpaceae5 (580spp.),Pandanaceae(700spp.),and
endemicNeotropicalfamilieshave over
Nepenthaceae(68 spp.). Six essentially
a hundredspecies, and two-Bromeliaceaeand Cactaceae-have about 2,000
5The discoveryofa primitive in SouthAmerica(Maguire& Ashton,1977)does not

dipterocarp
of "essentiallyendemic"used here.
bythedefinition
changethestatusofthefamily

TABLE 1. EndemicNeotropicalfamilies
(insomecases witha singleAfrican
speciesora single
monotypic
or ditypicAfrican
genuspresumably recently
arrivedbylong-distance
dispersal).
1) Tropicalforest taxa*
Bromeliaceae(46 genera/2, 108species(plus 1 sp. in Africa))
Caricaceae(3/29(plusone ditypicAfrican genus))
Caryocaraceae(2/24)
Cyclanthaceae(11/178)
Dialypetalanthaceae(1/1)
Duckeodendraceae (1/1)
Humiriaceae(8/46(plus 1 sp. in Africa))
Lacistemmaceae (2/14)(includedin Flacourtiaceae in FloraNeotropica)
Marcgraviaceae(4/125)
Quiinaceae(4/53)
Rapateaceae(15/79(plus 1 monotypic Africangenus))
Vochysiaceae(7/182(plus 1 ditypicAfrican genus))
Trigoniaceae/Trigonia(1/24)
Bignoniaceae/Bignonieae (46/359)
Lecythidaceae/Lecythidoideae (300)
2) Taxa of dryandlor Andean parts of South America (a few reachingNorthAmerica)
Brunelliaceae
(1/51)
Calyceraceae(4/46)
Columelliaceae(1/4)
Gomortegaceae (temp.)(1/1)
Malesherbiaceae (1/27)
Myzodendraceae (temp.)(1/11)
Nolanaceae(1/18)
Tovariaceae(1/2)(includedin Capparidaceaein FloraofPanama)
Tropaeolaceae(2/92)
3) Taxa of dryparts of tropicalNorthAmerica
Crossosomataceae (1/4)
Fouqueriaceae(2/8)
Garryaceae(1/18)
Lennoaceae(3/8)(1 rarespecieson thedryCaribbeancoastofnorthern Colombia)
Theophrastaceae(5/110)(a fewspeciesofJacquiniaand Clavija in SouthAmerica)
4) Taxa of drytropicallsubtropical
parts of both continents
Cactaceae(62/2,000) (a widespread speciesalso in Africaand Ceylonwithat least 1 derivative
in
Madagascar)
Julianaceae(2/5)(butprobablyan artificial group)
Koeberliniaceae(1/1)
Krameriaceae (1/15)
Loasaceae (12/266 (plus 1 in Africa))(severalsmallgenerain MexicoandSW U.S.)
Martyniaceae (3/13)
5) Miscellaneous (aquatics or semiaquatics and Guayanan edaphic specialists)
Cannaceae(1/55)
Cyrillaceae(3/13)(?) (in Europein Eocene; mostlyWestIndian)
Mayacaceae(1/9(also 1 in Africa))
Saccifoliaceae(1/1)
Sarraceniaceae(?) (3/17)(only1 genus(6 spp.) in tropics)
Tepuianthaceae (1/5)
Thurniaceae (1/3)
* I havefollowedtraditionalfamilial limits.Rhabdodendraceae (2 species)was treatedas a dis-
tinctfamily in FloraNeotropica.The otherfamilies listedbyPrance(1978)as endemicare segregates
(e.g., Peridiscaceae),erroneouslylisted(Pontederiaceae), or both(Heliconiaceae).

species each. Unlike most of theirPalaeotropical counterparts,manyof the Neo-

familiesare ecologicallyimportant.
tropical-restricted Bromeliaceaeis one ofthe
majorepiphytefamilies;Cactaceae dominatemanydryregions.Familieslike
Caryocaraceae,Humiriaceae,and Vochysiaceaeare important canopytreesof
lowlandtropicalforests.To thesemaybe added such specioseendemicgroups
as tribeBignonieaeof Bignoniaceae(Gentry,1980a),whichis thepredominant
groupof Neotropicallianas; subfamily Lecythidoideae(Lecythidaceae;Prance
& Mori, 1979),whichis one of the predominant taxa of canopytreesof Ama-
zonianforests;or subtribes Pleurothallidinae,
Maxillarinae,and Oncidinaeofthe
Orchidaceae,whichtogethercomprisenearly5,000species of Neotropicalepi-
phytes(Dodson,pers.comm.).
It is noteworthythatmanyoftheendemicfamilieslistedin Table 1 are taxa
ofdry,moreor less subtropical habitats.The onlyendemictropicalNorthAmer-
ican families-Crossosomataceae, Fouqueriaceae,Garryaceae,Koeberliniaceae
(disjunctto Paraguay),and Lennoaceae (also local in extremenorthern Colom-
bia)-are dryarea specialists.Although endemic,basicallySouthAmericanfam-
iliesare morediversifiedecologically,manyofthem,including Malesherbiaceae,
Nolanaceae, and amphicontinental Cactaceae, Martyniaceae, Julianaceae,Kra-
meriaceae,and Loasaceae are predominantly plantsofdryareas.
Although theseendemic taxa an
make appreciablecontribution to Neotropical
speciesrichness, thequestionofwhyfamilies thatevolvedintheNeotropicshave
speciatedmoreprofusely thantheirPalaeotropicalequivalentshas notyetbeen
addressed.
LAURASIAN TAXA
The phytogeographical significanceofthenewdevelopments inplatetectonics

was firstbrought intofocusbyRavenandAxelrod(1974).One ofthemajorthrusts
oftheRaven-Axelrod was thefundamental
synthesis differencebetweenthefloras
of the northern or Laurasiancontinents and thoseof thenow widelyseparated
southerncontinents thatwere clusteredtogetherat the timeof originof the
angiosperms. Evenbeforeplatetectonicsbecamegenerally accepted,ithadbeen
realizedthatthe highaltitudeSouthAmericanflorawas largelyderivedfrom
northtemperatesourcesand thatthe Panamanianisthmushad been a critical
barrierto the southward migrationof manynorthern taxa. One ofthecontribu-
tionsoftheRavenandAxelrodsynthesis was pointingouttheremarkable degree
to whichmanyplantfamilies, eventhosesharedbybothNorthand SouthAmer-
ica, can be unambiguously referredeitherto theGondwanaland or to theLaur-
asian florason accountoftheirgeneraldistributionalpatternsand thefossilrec-
ord. They listed51 basicallyGondwanaland plantfamilies(or equivalentunits)
thatprobablyspreadfromSouthAmericato NorthAmericain themiddleto late
Cenozoic as the Isthmianbarrierdecreased,and an additional54 families(or
equivalents)whoseoriginalpresencein NorthAmericawas probablyolderbut
whosemajorpresencethereprobablyresultedfrommid-to late-Cenozoicmigra-
tionfromSouthAmerica.Similarly a listof29 Laurasianfamilies(or equivalents)
whose majorarrivalin SouthAmericaprobablycoincidedwithbuildingof the
Isthmusof Panama in late Miocene to Pliocene was suggested,nine of them
qualifiedas perhapsalreadyhavinghad a priorpresencein SouthAmerica;11

TABLE 2. Laurasian elements of the Neotropical flora. Numbers indicate Neotropical genera
withknownspecies numbers/species in those genera(+ Neotropicalgeneraforwhichspecies estimates
are unavailable/totalspecies in those genera).
tAceraceae 1/5 (Krameriaceae) 1/15

Actinidiaceae 1/80 Labiatae (?) 14/489 (+13/1,594)
*Aquifoliaceae 1/150 tLennoaceae 4/8
Aristolochiaceae 3/182 Liliaceae 23/167 (+ 15/1,217)
tBalsaminaceae 1/2 *Lythraceae (?) 16/361
*Basellaceae 3/7 Magnoliaceae (+2/130)
Bataceae 1/1 Myricaceae (+ 1/35)
Berberidaceae 1/24 (+ 1/450) t(Nyssaceae) 1/1
Betulaceae 3/5 Oleaceae 8/52
Boraginaceae (?) 11/96 (+14/1,020) Orobanchaceae 1/2 (+1/140)
Buxaceae 3/42 Papaveraceae 1/1 (+3/120)
Callitrichaceae(?) (+1/25) tPinaceae 4/40
Caprifoliaceae 1/2 (+4/458) Plantaginaceae 1/1 (+ 1/265)
Caryophyllaceae 6/77 (+ 15/1,389) tPlatanaceae 1/7
*Celastraceae 16/102 (+4/361) Plumbaginaceae 2/7
*Chloranthaceae 1/40 Polemoniaceae 4/33 (+5/276)
Cistaceae (+3/120) Primulaceae 2/4 (+2/210)
Pyrolaceae 1/1 (+3/33)
Clethraceae 1/38 Rafflesiaceae 4/29
Cornaceae 2/8 (+ 1/6) Ranunculaceae 3/14 (+7/1,097)
Crassulaceae 3/25 (+4/515) *Rhamnaceae 23/168
tCrossosomataceae 1/4 Rosaceae 16/95 (+15/2,279)
Cruciferae 19/93 (+16/1,304) *Sabiaceae (?) 2/47
Cyrillaceae 2/13 Salicaceae 2/30
Droseraceae 1/20 Saxifragaceae 12/85 (+5/730)
Empetraceae 1/1 Scrophulariaceae 70/853 (+4/185)
Ephedraceae (?) (+1/40) Staphyleaceae 2/5 (+1/30)
Fagaceae 4/164 Styraceae 1/3 (+ 1/130)
tFouqueriaceae 2/8 Symplocaceae 1/160
t(Garryaceae) 1/18 *Theaceae 7/84 (+2/130)
Gentianaceae 19/494 Theophrastaceae 3/107
Geraniaceae 4/42 (+2/490) Typhaceae (+ 1/10)
tHamamelidaceae 3/5 *Ulmaceae 6/17 (+2/110)
Hippocastanaceae 2/3 Umbelliferae 48/480
Hydrophyllaceae 2/46 (+3/230) Valerianaceae 5/44 (+ 1/200)
tllliciaceae 1/1 *Vitaceae 2/3 (+3/505)
Juglandaceae 4/18
Total 416/5,229 (+ 157/15,834)
t = does not reach South America.
* = listed as movingfromSouth America to NorthAmerica by Raven and Axelrod (1974).
otherpredominantly Laurasianfamilieswere suggestedas probableearlierar-

rivalsin SouthAmerica.As thusinterpreted,themodernfloraoftropicalAmerica
is ofremarkably bipolarcomposition, withthegreatmajority
ofitsplantfamilies
havingclearlyLaurasianor clearlyGondwananaffinities.Fromthisperspective,
one mightanticipatethattheNeogenemixture oftwodifferentfloristic
elements
via the CentralAmericanisthmuscould have had a majoreffecton increasing
floristic perhapsalmostdoublingtheresultant
diversity, floraofeach ofthetwo
majorneotropicalregions.Althoughsuch floristic also occurredin
interchange
thePalaeotropics,it was less direct,interrupted
by largeexpansesof desertin
NorthAfricaanda persistent waterbarrierbetweentheSundaand Sahulshelves
in Australasia.
However,myanalysis(Tables2-6) suggeststhataddingtogether ofLaurasian

TABLE 3. Summaryof transectdata forplants over 1" dbh in a 1,000m2sample of lowland wet
forestat Los Tuxtlas, Veracruz (precipitationof 4,100 mm a year) (Gentry,in prep.).
No. Spp. No. Ind. No. Spp. No. Ind.

Gondwanan Families Palmae 4 113
Acanthaceae 1 1 Piperaceae 4 6
Anacardiaceae 1 2 Polygonaceae 2 2
Annonaceae 2 5 Rubiaceae 4 12
Apocynaceae 3 5 Sapindaceae 4 5
Araceae 3 3 Sapotaceae 3 8
Araliaceae 1 2 Solanaceae 3 5
Bignoniaceae 4 9 Tiliaceae 2 3
Bombacaceae 1 1 Urticaceaea 3 22
Capparidaceae 1 2 Violaceae 3 10
Caricaceae 1 2 Total 97 321
Compositae 2 3
Convolvulaceae 1 1 Laurasian Families
Dilleniaceae 2 7 Clsrca
Euphorbiaceae 4 10 Celastraceae 2 3
Rhamniaceae 1 2
Flacourtiaceae 2 2
S laceae 1 2
Hippocrateaceae 2 8
Lauraceae 5 9 Ulmaceae 1 1
Leguminosae 7 12 Total 5 8
Malpighiaceae 4 7 Unassigned Families
Meispermaceae 1 1 Malvaceaeb
Monimiaceae 1 3 Verbenaceae 2 2
Moraceae 7 24 indet. (cf. Fraxinus) 1 1
Musaceae 1 7 Total 4 4
Myrtaceae 3 3
Total: 106 species, including29 liana species. For a similar1,000m2sample in a South American
or southernCentral Americanforestwith4,100 mm of rain 170 species over 2.5 cm diam. would be
expected (based on the regressionof species numbersversus precipitationcalculated fromthe 19
sites of Gentry, 1981, 1982, and in prep.). The Veracruz diversityis significantly lower than the
expected inner tropicalvalue (R. Perozzi, pers. comm.). Note the overwhelmingpreponderanceof
Gondwana-derivedfamilies.
Assignmentas "Gondwanan" tentative.
a
The Los Tuxtlas genus is Robinsonella, an exclusively Central American (= tropical Laura-
b
sian?) genus, and the familymay be fundamentallyLaurasian in origin.
and Gondwanan elementshas not greatlyincreased overall Neotropical floristic

diversity.There are over 10 times as many Gondwanan-derivedas Laurasian-
derived Neotropical species. The northwardmigratingGondwanan taxa have so
overwhelmedthe correspondingsouthwardmigratingLaurasian taxa numerically
that the latter's contributionsto the total Neotropical florahave generallybeen
relativelyinsignificant.This patternis especiallyprevalentin thetropicallowlands
ofCentralAmerica,whichmustonce have been populatedby a tropicalLaurasian
floristicequivalentofthe endemicCentralAmericanherpetofauna(Savage, 1966),
mammalianfauna (Patterson& Pascual, 1972) or avifauna (e.g., Cracraft,1973)
(see Raven & Axelrod, 1974: 625-626). Graham (1976, 1982) has shown that a
few South Americantaxa such as Dichapetalum, Casearia, Laetia, Symphonia,
Gustavia, and Byttneriahad alreadyreached Veracruz, Mexico by Miocene times.
Yet the Paraje Solo palynoflorawas dominated by temperateNorth American

TABLE 4. Amazonian-centereaGondwanan families.NumbersindicateNeotropicalgenera with

known species numbers/neotropical species in those genera (+ Neotropical genera forwhich species
estimatesare unavailable/totalspecies in those genera).
Anacardiaceae (?) 15/133 (+2/257) (Lacistemaceae) 2/14

Annonaceae 28/555 (+3/250) Lauraceae 11/700 (+4/870)
Apocynaceae 64/687 (+2/125) Lecythidaceae 11/275
Bignoniaceae 72/631 Leguminosae 216/2,980 (+48/8,189)
Bixaceae 1/5 Loganiaceae 12/136 (+2/106)
Bombacaceae 20/187 Malpighiaceae 44/801
Burseraceae (?) 5/102 (+2/120) Meliaceae 8/125
Caryocaraceae 2/24 Menispermaceae 17/142 (+ 1/30)
Chrysobalanaceae 8/334 Moraceae 23/408
Cochlospermaceae 2/8 Myristicaceae 5/81
Combretaceae 7/97 Ochnaceae 9/67 (+ 1/300)
Connaraceae 4/57 (+ 1/100) Olacaceae 13/87
Convolvulaceae 21/1,000 Palmae 52/1,110 (+3/42)
(Dialypetalanthaceae) 1/1 Polygalaceae 6/183 (+3/630)
Dichapetalaceae 3/43 Quiinaceae 4/53
Dilleniaceae 5/60 Rhizophoraceae 5/24
(Duckeodendraceae) 1/1 Sapindaceae 27/438 (+5/490)
Ebenaceae 2/82 Sapotaceae 9/208 (+3/234)
Elaeocarpaceae 4/7 (+2/125) Simaroubaceae 11/106
Euphorbiaceae 92/2,607 Sterculiaceae (?) 14/293 (+2/360)
Flacourtiaceae 28/267 Tiliaceae 20/139
Gnetaceae 1/6 Trigoniaceae 1/24
Hernandiaceae 3/22 Turneraceae (?) 1/60 (+2/26)
Hippocrateaceae 12/114 Violaceae 11/98 (+2/650)
Humiriaceae 8/46 Vochysiaceae 7/182
Icacinaceae 13/56 Total 961/15,866 (+88/12,904)
elements.Similarlyan Oligocenesite in PuertoRico (Graham& Jarzen,1969)

was characterized by thepresence,althoughat reducedlevels,of severalofthe
same northtemperate generathattodayare disjunctin themidaltitude"bosque
caducifolia"ofMexico:Liquidambar,Fagus, Nyssa as wellas suchotherLau-
rasiantaxa as Myrica,Engelhardtia,and Hauya. These taxa led Grahamand
Jarzento emphasizeprior,moredirectmigration betweenMexicoandtheGreater
Antilles.Nevertheless a numberoftropicalSouthAmericantaxa thatmusthave
arrivedoverwaterwerepresent.
As the Isthmianconnectionclosed, additionalSouthAmericantaxa moved
northto completely dominatethe CentralAmericanlowlands.Most of thisin-
vasion has been so recentthateven at the specificlevel therehas been little
differentiation.
Thus,in groupslike tribesTecomeae and Bignonieaeof Bigno-
niaceae,virtually all of the speciesthatreachnorthern CentralAmericaare in-
distinguishablefromSouthAmericantaxa (compareGentry,1982cwithGentry,
1973).Thereare onlyone speciesofTecomeaeand sevenofBignonieaein Gua-
temalathatare not also in Colombiaand Venezuela. Perhapsthisnorthward
migration is-stilltakingplace. At anyrate,thereis a clearnorthward
decreasein
the numberof species of manyNeotropicalfamilies(Gentry,1982a).It seems
likelythat,in general,individualtropicallowlandforestsin northern Central
Americamaybe less diversethantheirsouthernequivalents,as suggestedby
Sarukhan(1968).Toledo (1982)has shownthatwithinMexicotreespeciesrich-

TABLE 5. Andean-centeredGondwanaland groups. Numbers indicate neotropicalgenera with

knownspecies numbers/Neotropical species in those genera (+ Neotropical genera forwhich species
estimatesare unavailable/totalspecies in those genera).
Sa. NorthernAndes
Acanthaceae 61/1,493 Loranthaceae 16/592 (+ 1/15)
Araceae 38/1,386 Marantaceae 10/270 (+ 1/11)
Araliaceae 5/197 (3/356) Marcgraviaceae 4/125
Balanophoraceae 7/15 Melastomataceae 85/3,153
Begoniaceae 1/600 Monimiaceae 8/246
Bromeliaceae 46/2,108 Musaceae 2/82
Brunelliaceae 1/51 Myrsinaceae 12/311 (+2/600)
Campanulaceae 9/568 (+7/712) Nyctaginaceae ? 27/277 (+3/160)
Cannaceae 1/55 Orchidaceae 306/8,266
Caricaceae 3/29 Oxalidaceae 1/8 (+2/870)
Columelliaceae 1/4 Passifloraceae 4/362
Compositae 502/3,864 (+87/7,202) Piperaceae 4/25 (+2/3,000)
Cyclanthaceae 11/178 Rubiaceae 147/2,906 (+21/2,545)
*Ericaceae 37/731 Tovariaceae 1/2
Gesneriaceae 40/917 Tropaeolaceae 2/92
Guttiferae 21/232 (+3/590) Urticaceae ? 7/88 (+6/653)
Haloragidaceae 1/1 (+3/58) Zingiberaceae 4/111
Total 1,425/29,345 (+ 141/16,772)
Sb. SouthernAndes/SouthTemperate
tAetoxicaceae 1/1 Loasaceae 12/266
tAuraucariaceae 1/2 tMalesherbiaceae 1/27
Calyceraceae 4/46 Myrtaceae 24/1,254 (+2/1,100)
Coriariaceae 1/1 tMyzodendraceae 1/11
Cunoniaceae 3/12 (+ 1/170) tNolanaceae 1/18
Cupressaceae 3/5 *Onagraceae 14/275
tEpacridaceae 1/1 Podocarpaceae 1/37
tEucryphaceae 1/1 Portulacaceae 5/8 (+5/422)
tFrankeniaceae 3/8 Proteaceae 3/92
tGomortegaceae 1/1 tRestionaceae 1/1
Hydnoraceae 1/6 Santalaceae 7/43
Iridaceae 34/188 (+ 1/100) Solanaceae 66/1,861 (+1/8)
Juncaceae 6/49 Winteraceae 1/1
tLardizabalaceae 2/3
Total 199/4,218 (+10/1,800)
t = does not reach CentralAmerica.

* = listed as movingfromNorth America to South America by Raven and Axelrod (1974).
ness of the lowlandtropicalrainforestdecreasesdramaticallynorthward. My

data froma 1,000m2sampleof richlowlandrainforestin Veracruzalso show
fewerspeciesthanwouldbe expectedin a simiilarvegetationfurther
south(Table
3). However,thispatternis shownonlybylowlandmoistforestspecies:Mexican
dryareascontrast inbeingverydiverse,eveninultimatelysouthern-derivedtaxa,
withmanyendemicspecies(cf.,Rzedowski,1978:75).
The relatively
depauperateconditionoflowlandCentralAmerican forestsmay
also have a muchmorerecentoriginand be due largelyto Pleistoceneclimatic
fluctuations.Whiledrought has been consideredthemajoreffectofPleistocene
glaciationon thelowlandtropics,CentralAmerica,at themarginofthetropics,
mayhave been moreaffectedby the concomitant generalloweringof thetem-
perature;manysensitiveinnertropicaltaxa mayhave been eliminated or "con-

TABLE 6. Miscellaneous taxa. Numbersindicateneotropical

generawithknownspeciesnum-
bers/Neotropicalspeciesin thosegenera(+ Neotropical
generaforwhichspeciesestimates
are un-
available/total
speciesin thosegenera).
Guayana-centeredgroups Ceratophyllaceae 1/2

Burmanniaceae 13/51 Chenopodiaceae 10/34 (2/14)
Dipterocarpaceae 1/1 Commelinaceae 17/163
Mayacaceae? 1/9 Cucurbitaceae 55/311
Podostemataceae 19/151 Cyperaceae 8/88 (+ 18/3,738)
Rapateaceae? 15/79 Dioscoreaceae 1/15 (+1/600)
Saccifoliaceae 1/1 Elatinaceae (+2/45)
Sarraceniaceae? 1/6 Eriocaulaceae 12/868
Tepuianthaceae 1/1 Goodeniaceae 1/1
Triuridaceae 4/12 Gramineae 127/838 (+80/4,692)
Total 56/311 Haemodoraceae 1/2
Hydrocharitaceae 8/20
Dry-area Gondwanan groups Juncaginaceae 2/5
Cactaceae 60/2,000 Lemnaceae 4/12
Capparidaceae 10/40 (+4/416) Lentibulariaceae 3/116
Erythroxylaceae 1/180 Linaceae ? 3/20 (+1/230)
Koeberliniaceae 1/1 Malvaceae 50/860
Martyniaceae 3/13 Myoporaceae 1/1
Velloziaceae 4/229 Najadaceae 1/8
Zygophyllaceae 12/62 Nymphaeaceae 5/27
Total 91/2,525 (+4/416) Phytolaccaceae 13/78 (+2/38)
Polygonaceae 10/203 (+5/724)
Unassigned Pontederiaceae 4/21
Aizoaceae 2/4 (+4/90) Potamogetonaceae 5/42
Alismataceae 2/61 Rutaceae 36/233 (+4/313)
Amaranthaceae 7/188 (+7/535) Taccaceae (+ 1/30)
Amaryllidaceae 26/799 (+3/210) Thymelaeaceae 7/71 (+ 1/10)
Asclepiadaceae 46/932 (+3/280) Verbenaceae 40/1,143
Butomaceae 2/7 Xyridaceae 2/21 (+ 1/250)
Canellaceae 3/11 Total 516/7,206 (+135/11,799)
finedsouthward"bytheslightly lowertemperatures duringglacialadvances(Ax-

elrod,pers. comm.).This mayhave been theultimatefateofthetropicalNorth
Americanflorathatis knownto have inhabitedeven muchof the southernand
centralUnitedStatesduringtheEocene and wouldbe consistent withsuchpat-
ternsas themodemdiversity of Sabiaceae, knownto have been widespreadin
TertiaryNorthAmericantropicalfloras,whichis greaterin Panamaand Costa
Rica thanit is in northernCentralAmerica(Gentry,1981).
The northward movement of lowlandtropicalGondwananelementshas had
no significantcounterpart
ofsouthward movingtropicalLaurasiantaxa.The most
clearlyLaurasianfamilies to havenoteworthy complements ofspeciesinlowland
SouthAmericanforestsare Aristolochiaceae and Vitaceae,each witha single
vinegenuswithnumerousspeciesin SouthAmerica(Aristolochia, Cissus); not
surprisingly,botharealso wellrepresentedintheWestIndies.Fourothertropical
lowlandfamiliesof probableLaurasianderivationare characterized by affinity
fordryareas and a strongrepresentation bothin theWestIndiesand in northern
CentralAmerica.Threeof these-Buxaceae, Boraginaceae,and Rhamnaceae-
are proportionately betterrepresentedin temperate NorthAmericathanin the
Neotropics.In theirdryarea preference, these groupsare reminiscent of the

endemictropicalLaurasianfamiliespreviously noted.However,representatives
of all ofthesefamilieshave penetrated intolowlandSouthAmerica.In thecase
of Buxaceae, penetration of SouthAmericais minimal(contrary to thedistribu-
tion shownin Heywood, 1978) and restricted to a few limestoneoutcropsin
northern Venezuela(Stylocerasgoes southin theAndesbutmaynotbe closely
related;Gentry& Foster,1981).Theophrastaceae has morerecordedspeciesin
Peruthanfarther northbutis a morepredominant vegetationalelementand has
greatergenericdiversity in CentralAmericaand the WestIndies. One genus,
Clavija, whichlinksTheophrastaceae to Myrsinaceae, occursinlowlandtropical
forestsbutis poorlyrepresented inAmazoniaand mayhaveonlya singlespecies
reachingcoastal Brazil.Onlytwo genera(Cordia, Tournefortia) of thetwenty-
fourgeneraof Boraginaceaethatreach the Neotropicspenetratethe lowland
tropicalforeststo any extent.AlthoughRhamnaceaewere listedas basically
GondwananbyRavenandAxelrod(1974),thepatternshownbyRhamnus(John-
ston & Johnston,1978) seems typicalof the groupand pointsto a northern
ancestry.As in Boraginaceae,thepenetration of thefamilyintolowlandNeo-
tropicalforestsis minimal(monotypic Ampelozizyphus and a few species of
Gouania and Colubrina),althoughitis betterrepresented in drierpartsofSouth
America.
WhilethelowlandtropicalSouthAmericanflorawouldbe almostimpercep-
tiblychangedifall oftheseputatively tropicalLaurasiangroups(a totalofperhaps
a fewhundredspeciesin all oflowlandtropicalSouthAmerica)wereeliminated,
Laurasiantaxa are muchmoreimportant in Neotropicalmontanefloras.In fact,
thereseemsto be a basic dichotomy betweentheLaurasian-derived uplandand
Gondwanan-derived lowlandneotropical floras.In CentralAmericanuplandfor-
ests Laurasianelementsclearlypredominate ecologicallywithfamilieslike Pi-
naceae, Fagaceae, Juglandaceae, Magnoliaceae,Theaceae, and Ulmaceae espe-
ciallyimportant as canopymembersof the temperatemontaneforests.These
northern taxagradually decreasesouthward so thatfamilies likeHamamelidaceae
and Pinaceae do notcrosstheRio San Juanlowlandsand are notpresentsouth
of northern Nicaragua,whileGarryaceaeand manyimportant generaof other
families(e.g., Ulmus,Celastrus,mostJuglandaceae) reachonlyuplandPanama.
Even in SouthAmerica,Laurasianelementstendto prevailin montanefor-
ests,ecologically, ifnotalwaysin numbersofspecies.Manyofthesespeciesare
wind-pollinated and thusespeciallywell represented in thefossilrecord.Con-
sequentlywe maybe reasonablyconfident thatthepalynological documentation
of theirrecentarrivalin SouthAmericais meaningful. Such knowledgeof the
historyof Andeanforestsreliesalmosttotallyon theworkof van derHammen
andhisassociates(summary invanderHammen,1974).Thefirst montane elements
toarriveatthePalynological sitesintheCordillera OrientalattheSabanade Bogota
wereHedyosmum andMyrica,as theCordillera was uplifted duringthePliocene.
By the beginning of the Pleistocene,the principalupheavalof the regionwas
completed.DuringtheearliestPleistoceneglacialadvancethepalynoflora ofthis
regionsuggesteda primitive and depauperateparamovegetation includingsuch
ultimately northern-derived elementsas Aragoa (Scrophulariaceae), Hypericum
(Hypericaceae),Umbelliferae, Plantago,Polylepis(Rosaceae; perhapssouthern),
Valeriana,and Ranunculaceae.DuringthePleistocenethepalynoflora fluctuated

TROPICAL AMERICA Ldyer 1ft Md) Fkxd NewIr opct ki

b rr no 3
FIGURE1. TypicalAmazon-centered distributionof a taxonof canopytrees,Moraceaetribe

withspeciesdiversity
Olmedieae.Tribaldistribution isohyetsplottedfromthedata of Berg(1972:
ofspeciesin (wetter)westernAmazoniain thearea oftheColombia/
fig.1). Note theconcentration
Brazil/Peru Additional
frontier. in poorlyknownnorthern
collecting AmazonianPeruand adjacent
Colombiawillprobably extendthehighdiversityregionwestward.
withthechanging altitudinalzonationofthevegetation broughtaboutby climatic

changesbetweenglacialand interglacial periods.Boththeparamoand montane
forestflorasweregraduallyenrichedduringthePleistocene.In thelowerPleis-
tocene,such additionalnorthern elementsas Geranium,Gentiana,Lysipomia,
Juglans,and (perhapssouthern)Urticaceaeappearin thepollenrecord,along
withsoutherntaxa like Gunneraand phytogeographically problematicalStylo-
ceras (see Gentry& Foster,1981).Alnusfirstarrivedat the end of thelower
Pleistoceneand becamedominantduringthe middlePleistocene.Quercusfirst
appearedapproximately 250,000yearsago at theend of theMiddlePleistocene
and thereafterincreasedprogressively in importance.Althoughsome southern
taxa like Weinmannia also arrived in the Cordillera duringthePleisto-
Oriental
cene, northern elementsprevailed and the presentnorthernAndean forestsare
stilldominatedby Laurasian taxa.
Even todaysuchnorthern Betulaceae,
familiesas Myricaceae,Juglandaceae,

O~~A>D _ _I_A
l _ 3C__
_
0-Lt
97 ; nb f i r i )
Fagaceae, Magnoliaceae,Berberidaceae,Hippocastanaceac,Cyrillaceac,Cle-
thraceae,Cornaceae,Oleaceae, and Caprifoliaceac are presentin tropicalSouth
Americaalmostentirely in the uplandAndes. WithintheAndes,thereis a de-
crease in representation of thesefamiliesfarthersouth.For example,Quercus,
theabsolutedominant ofmostColombianlowermontaneforests,does notoccur
in Ecuador. Otherwoodyfamilieslike Salicaceae, Ulmaceae,Theaceae, Celas-
traceae,Aquifoliaceae,Sabiaceae, and Staphyleaceae,have one or two wide-
spreadspecies(or genera)thathavebecomewidespreadin thetropicallowlands
(respectively: Salix humboldtiana,Trema mnicrantha and Celtis iguanea, Tern-
stroemia, Gouepia, hlexinundata, Ophiocaryon,Turpiniaoccidentalis). Interest-
ingly,thelowlandrepresentatives of suchtaxa are oftenrestrictedin Amazonia
to ecologicallyimpoverished extremesitessuchas seasonallyinundatedstream-
sides (Salix, hlex, Ophiocaryon duckei, Ampelozizyphus(Rhamnaceae)),white
sand substrates (manyTernstroemiaand Ophiocaryon), or secondgrowth(Tre-
ma, Celtis).Similarly, theonlyAmazonianspeciesof southtemperate Podocar-
pus is restricted to whitesand (Gentryet al. 2887] (MO) fromnear Iquitos,
apparently an undescribed species).
Predominantly herbaceousLaurasianfamilieshave a greatertendencyto be

C
SUEZ ~~~~Guzmania
.
\ Vriesea - - ~ - h
C- CO- tXCT-00 S.\iltondito ,i la
Tillondsia
_ ~~~~~~~~~~~Vriesea
~ ~~~~~~~~~
~~Catopsis ~
>
~ rise
_ t \
Vriesea
vet~~~~~~~~~~~~~~
Tilan'i ~~\s
I~~~~~~~~~~~
a f t ~~~~~~Tillandsia
distribution
orAndean-centered
FIGURE 3. Extra-Amazonian tax-
epiphytic
ofa predominantly
on (Bromeliaceae,
subfamily Smith& Downs,1977:fig.213).
Tillandsioideae;

576 ANNALSOF THE MISSOURIBOTANICAL
GARDEN [VOL.69
FIGURE 4. Neotropical regions.

phytogeographic 1. MexicoandCentral
America.
2. WestIn-
dies. 3. Northern
Venezuela-Colombia. 4. Northern Andes.5. SouthernAndes.6. Amazonia(western
borderdefined by500mcontour). 7. GuayanaHighlands (over500in). 8. Guianasubregion(included
as partofAmazoniaexceptforspeciesnotfoundelsewhereinAmazonia).9. Cerradoandassociated
dryareas. 10. CoastalBrazil.Smallnumbers indicatepercentof monographed speciesoccurringin
or endemicto each region.T = Percentoftotalsampleof8,117recently monographed speciesoc-
cutrngin thatregion.C = Canopytreesand lianas:percentofthespeciesoftaxacharacterized by
habitoccurring in each region(percentofregion'sspeciesofthathabitgroupwhichare endemicto
theregioninparentheses). E = Epiphytes andpalmettos: percentofthespeciesoftaxacharacterized
by habitoccurring in eachregion(percentofregion'sspeciesofthathabitgroupwhichare endemic
to theregionin parentheses).See Tables7, 8, 9 forcompletedata.
weedyand theirpatterns are notso wellmarked.Nevertheless, exceptfora few

weeds,familieslikeCrassulaceae,Caryophyllaceae,Ranunculaceae,Cruciferae,
Saxifragaceae,Rosaceae (s.s.), Plumbaginaceae,
Geraniaceae,Callitrichaceae,
Balsaminaceae,Umbelliferae, Primulaceae,Gentianaceae,Polemoniaceae,Hy-
drophyllaceae,
Scrophulariaceae, Orobanchaceae,and Plantaginaceaeare much
betterrepresentedin themontaneNeotropicsthanin thelowlands.
For bothtreesand herbsthereis a strongdichotomy betweenthenoticeable
presenceof Laurasiantaxa in montaneforestsand theirvirtualabsence in the
lowlands.

MostoftheLaurasiantaxa,especiallythewoodyones,have speciatedrather
littleintheAndes(see Gentry,1982a).Presumably inparttheirrecent
thisreflects
listof72 Laurasian-derived
arrival.As a result,theimpressive Neotropicalfam-
ilies in Table 2 accounts for a very small percentage (<10%o) of the total Neo-
noneofthatofthelowlandtropics.
tropicalfloraand virtually
GONDWANAN TAXA
The majorcomponents of the Neotropicalfloraare Gondwanaland-derived
groups(Tables4-6). However,thereis a fundamental phytogeographic difference
withinthisgroup,fortheseautochthonous SouthAmericanfamiliesfallconsis-
tentlyintotwomajordistributional groups.Manyof thesefamiliesare centered
in Amazonia(Table 4; Fig. 1) and maybe referred to as Amazonian-centered.
Mostoftheremainder arefundamentally andcontrastingly extra-Amazonian with
verypoorrepresentation in Amazoniaand usuallyhave theirdistributional cen-
tersintheAndes,especiallythenorthern Andes(Table 5; Figs. 2, 3). Thesetaxa
mayconveniently be termedAndean-centered taxa although theirmaindiversity
is reachedinthelowlandsnearthebase ofthemountains andinmiddleelevation
cloudforestsratherthanin thehighmountains themselves.
I firstbecameaware of thesetwo striking patternsin thefield.In orderto
documentthem,I extractedand compileddistributional dataforrecently mono-
graphedNeotropicalfamiliesand largegenera,beginning withtheFloraNeotro-
pica monographs, butalso including as manyotherFloraNeotropica-caliber re-
visionsof majortaxa as I couldfindin majorsystematic botanyjournalsforthe
last tento twentyyears.This yieldeda data set of 8,117monographed species.
Subdividing theNeotropicsintotheninephytogeographic regionsshownin Fig-
ure4, and counting thenumberofmonographed speciesoccurring in each region
providedthe data summarized in Tables 7 and 8. The greatmajority(71% or
almost5,800species)of monographed speciesbelongto familiesthatare either
clearlyAmazonian-centered (3,052 spp.) or clearlynorthern-Andean-centered
(2,715spp.). The component familiesof each of thesemajorphytogeographical
groupsnotonlyshowstrikingly concordant distributional
patterns butalso amaz-
ingconsistency ecologically.The Amazonian-centered taxa are overwhelmingly
canopytreesandlianas;theextra-Amazonian ones are chieflyshrubs,epiphytes,
and palmettos.
AMAZONIAN-CENTERED TAXA
The Amazonian-centered taxa include38%oof the totaldata set of mono-

graphedspecies. These familieshave an averageof almosthalf(44%) of their
speciesin Amazonia.Virtually canopytrees
all ofthesetaxa are predominantly
and lianas;moreover, almostall canopytreesand lianasoftheNeotropicallow-
landsbelongto theseAmazonian-centered groups.Thesetaxaare conspicuously
under-represented in theAndes;on theaverageonly12%oftheirspeciesoccur
inthenorthern Andeanregionand only5% intheSouthern Andes.Theyare well
represented in thecoastalBrazilregion(16% oftheirspecies),whichconstitutes
a distinctsecondarycenterformostofthem;however,nota singleone ofthese
monographed majortaxa has as manyspeciesin coastalBrazilas in Amazonia,

TABLE 7. Numberofspeciesandendemism
inmonographed
taxabypredominant
habitgroupandg
speciesforeachregioninparentheses.
Central
Cerrado America
Coastal & North. South. &
Habit' Amazonia Brazil Caatinga Andes Andes Mexico
Canopy trees and lianas 1,334(1,072) 482 (322) 373 (272) 373 (154) 144 (85) 453 (189)
Epiphytes
andpalmettos 292(205) 498 (426) 184(140) 723(481) 469(231) 607(443)
Herbsand shrubs 60 (16) 140(76) 167(103) 128(51) 437(231) 559 (319)
Vines3 75 (40) 74 (40) 45 (18) 126(73) 78 (40) 132(65)
Montane trees4 7 (5) 12 (8) 3 (1) 71 (47) 57 (39) 98 (84)
Aridarea trees 2 (1) 17 (12) 4 (1) 47 (37) 37 (30)
Miscellaneous5 61(52) 80 (67) 143(130) 29 (12) 31(20) 48 (35)
Total (8,117) 1,829 (1,390) 1,288 (940) 932 (676) 1,454 (819) 1,263 (683) 1,934(1,165)
Percentofspecies *0
occurringin region 23% 16% 11% 18% 16% 24%
% Endemism6
(total=
6,567/8,117= 81%) 76% 73% 73% 56% 54% 60%o
Totalendemism7 17% 12% 8% 10%70 8% 14%
'Predominant habitofmonographed taxon.
2
FiguresforGuianassubregion includeonlythosespeciesthatoccurin thelowlandGuianasbutnot
Distribution
primarilyreflects
patternofsinglelargefamily-Passifloraceae (363spp.).
4 Betulaceae,Brunelliaceae,Clethraceae,Juglandaceae,Polylepis,Podocarpaceae,Rhamnus,Colu
manyMexicandesertshrubs.
5 Parasites,
aquatics,Velloziaceae,Proteaceae.
6 Endemic speciesin regionas percentoftotalmonographed speciesin thatregion.
7 Endemicspeciesin regionas percent oftotalspeciesin all monographed taxa.

TABLE ofmonographed
8. Percentdistribution distrib
habitgroupandgeographical
taxabypredominant
Central
Cerrado Amer. N. V
Coast. & North. South. & zue
Amaz. Braz. Caatinga Andes Andes Mexico Colo
Total Percentof
Habit' Spp. Species3
Canopytreesandlianas 3,052 44 16 12 12 5 15
andpalmettos
Epiphytes 2,715 11 18 7 27 17 22
Herbsand shrubs 1,034 6 14 16 12 42 54 1
Vines4 459 16 16 10 27 17 29
Montanetrees5 256 3. 5 1 28 27 38
Arid area trees 105 2 16 4 45 35
Others(parasities,
aquatics,Velloz.,
Proteac.) 496 12 16 29 6 6 10
Total 8,117
I Predominant habitofmonographed taxon.
2Figures forGuianassubregion includeonlythosespeciesthatoccurin thelowlandGuianasbutnotalso i
3Percents totalto morethan100%osincewidespread speciesare countedin eachregionin whichtheyoccu
patterns
reflects
4Distributionprimarily ofsinglelargefamily-Passifloraceae(363spp.).
Clethraceae,
5Betulaceae, Brunelliaceae, Polylepis,Podocarpaceae,
Juglandaceae, Rhamnus,Colubrina

not even Dilleniaceae,whose strongrepresentation in the coastal regionwas

emphasizedby Kubitzki(1975). Manyof theAmazonian-centered familieshave
derivativespeciesin thecerradoand associateddryareas oftheBrazilianshield
(12% of the species); in almosteverycase the cerradospecies are shrubsor
subshrubsin taxa thatare otherwisetreesor lianas.These groupsare markedly
poor in the WestIndies(10%oof the species on the average,witha dispropor-
tionatepartofthattotaldue to theevolutionary explosionof a singleotherwise
smallsectionof Tabebuia).
The Amazonian-centered taxaarepoorlyrepresented inCentralAmerica(only
15% oftheirspecies),whichis rathersurprising sincetheymakeup nearlyall of
the moistand wet forestcanopyof the CentralAmericanlowlands,just as in
Amazonia.Most of thespeciesof thesetaxa thatdo reachCentralAmericaare
notendemic;rather,theyare mostlythosefewAmazonianspeciesthathappen
to have unusuallywideranges.Thus,only6%oofthe3,000monographed species
ofAmazonian-centered taxaare endemicto CentralAmerica.Thiscontrasts with
80%oendemismin Amazonia,where35% of all the species of these taxa are
endemic.Within CentralAmericathereis a markeddecreaseintherepresentation
of Amazonian-centered taxa fromsouthto north.Most of thesefamilieshave
severalspeciesreaching easternPanama,noticeably fewerreachingwesternPan-
ama and CostaRica,andnonecrossingtheHoldridgesystemtropical/subtropical
delimitation at 12'N latitudein Nicaragua(exactlythe same latitudeas the di-
versity-reducing Isthmusof Kra in Malaysia!).Those Amazonian-centered fam-
iliesthatdo extendfarther intoCentralAmericatypically have onlyone or two
species northof Nicaragua(see Gentry,1982a,forspecificexamplesof these
patterns).Nevertheless, therelativelyfewAmazonian-centered taxa thatreach
northern CentralAmericacontinueto constitute virtuallyall ofthelowlandforest
canopy(Table 3 and Gentry, in prep.).
An interesting subsidiary patternis shownin CentralAmericaby severalof
thesetaxa. Severalofthegroupshave a distinctsecondaryradiationin northern
CentralAmerica.A good exampleis providedby BignoniaceaewithtribeCres-
centieaehavingthreegeneraand35 speciesalmostexclusively inCentralAmerica
(Gentry,1979,1980a).Although derivedfromthefundamentally SouthAmerican
tribeTecomeae,Crescentieaeare so distinct fromthatgroupin such important
featuresas indehiscent fruitsandbat-pollinated flowersthattheyhave sometimes
beentreatedas a distinct family(see Gentry,1974a).ClearlyCrescentieaereflect
a longhistoryof differentiation in CentralAmericasubsequentto an initialcol-
onizationby SouthAmericanTecomeaestock.Yet mostotherCentralAmerican
Bignoniaceaeareundifferentiated fromtheirSouthAmericanprogenitors evenat
thespecieslevel.
It is temptingto thinkofsuchpatterns as reflectinga twopulsemigration: (1)
earlycolonizationby islandhoppingacross theproto-Antilles at theend of the
Cretaceouswithsubsequentmajordifferentiation and (2) a majormigration sub-
sequentto closing-of theIsthmusofPanamathatwas too recentto permitmuch
generic,or even specific,differentiation.
ANDEAN-CENTERED
TAXA
The second major Neotropical phytogeographicpattern,contrastinglyextra-
Amazonian, may be referredto convenientlyas Andean-centeredand is almost

themirror imageofthatshownbytheAmazonian-centered taxa.In thoseregions

whereAmazonian-centered taxa are wellrepresented,Andean-centered taxa are
poorlyrepresented and vice versa. Familieswiththispatternhave theirdistri-
butionalcentersinthenorthern Andes,whereovera fourth (27%) oftheirspecies
occur,and are also wellrepresented in thesouthern Andes(17% oftheirspecies)
(Table 8). These groupsare predominantly epiphytic
(Araceae,Araliaceae,Bro-
meliaceae,Cyclanthaceae,Ericaceae,Gesneriaceae,Guttiferae, Piperaceae/Pe-
peromia,Orchidaceae,etc.), understory shrubs(Acanthaceae,Caricaceae,Me-
lastomataceae, Monimiaceae,Myrsinaceae, Piperaceae,Rubiaceae,Solanaceae),
and coarse palmetto-type monocots(Musaceae, Marantaceae,Zingiberaceae).
These groupsare notonlyconspicuously under-representedin Amazonia(11%
of theirspecies),theyare also poorlyrepresented in the drycerrado-caatinga
region(7% of theirspecies). Like theAmazonian-centered group,theyare well
represented in thecoastalBrazilregion(18% oftheirspecies)and poorlyrepre-
sentedin northern Venezuelaand theWestIndies.
UnliketheirAmazonian-centered counterparts,theAndean-centered taxaare
verywell represented in CentralAmerica,especiallyCosta Rica and Panama,
where22% of theirspeciesoccur. SouthernCentralAmericais clearlya major
secondarycenterofspeciationformostofthesegroups.Although someofthese
groupsactuallyappearto have morespeciesin Costa Rica or Panamathaninthe
northern Andes,thismaybe mostlyan artifact ofthemuchpoorerfloristic data
base fromnorthwest SouthAmerica.In anyevent,thesegroupsaccountformost
of the incrediblefloristic diversityof the Choco region(Gentry,1982a). The
Andean-centered taxashowverypronounced endemism inCentralAmerica,with
73% of the CentralAmericanspeciesendemic.This is in strongcontrastto the
low (42%) CentralAmericanendemismof Amazonian-centered taxa (Table 9).
ClearlybothCentralAmericaand westernSouthAmericahavebeen majorevo-
lutionary centersforthesegroups.
Althoughrepresentation of thesetaxa is highestin mountainous phytogeo-
graphicregions,it shouldbe re-emphasized thathighspeciesdiversities do not
occurat highaltitudes butratherinthewetlowlandandpremontane cloudforests
alongthebase and lowerslopesofthemountains.
Thesetwodominant phytogeographic patterns-Amazonian-centered treesand
lianasand Andean-centered palmettos,shrubs,and epiphytes-together account
forthegreatmajority (71% ofmysample)ofNeotropicalplantspecies.Together
thesefamilies,all basicallyGondwanan,absolutelydominatethelowlandneo-
tropicalflora,bothin Centraland SouthAmerica.Thus any explanationof the
patterns ofevolutionary diversificationin thesetaxa willlargelyexplaintherich-
ness oftheNeotropicalflora.
DRY-AREA-CENTERED
TAXA
A fewsubsidiary distributional
patternsneed to be mentioned.One is thatof
taxa withdistributionalcentersin dryareas and poor representation both in
Amazoniaand themoistAndes. Threegood examplesare Capparidaceae,Cac-
taceae, and Zygophyllaceae.Amaranthaceae and possiblyChenopodiaceae,the
latteroftenspecializedforthehighlyalkalineconditionstypicalof deserts,are
also betterrepresentedin drythanin wetareas. Dry-area-centered
taxa tendto
be largelyshrubsand herbsalthoughsomewellknowntreegeneralikeProsopis

TABLE ofphytogeographic
9. Relativeendemism in eac
regions.Percentofthosespeciesoccurring
Ce
Total% Cerrado A
Endemic Coast. & North. South.
Habit' to I Reg. Amaz. Brazil Caatinga Andes Andes M
Canopytreesandlianas 81 80 67 73 41 59
andpalmettos
Epiphytes 81 70 86 76 67 49
Herbsand shrubs 82 27 54 62 40 53
Vines2 71 53 54 40 58 51
Montanetrees3 85 71 67 33 66 68
Arid area trees 84 - 50 71 25 79
'Predominant habitofmonographed taxon.

2Distribution patterns
reflects
primarily ofsinglelargefamily-Passifloraceae(363 spp.).
Juglandaceae,
Clethraceae,
3 Betulaceae,Brunelliaceae, Polylepis, Podocarpaceae,Rhamnus, Colu

and Bulnesia also show thispattern.These groupsare best represented in the

southern Andeanregion(42% oftheirspecies),hereincluding partofthemonte,
and the CentralAmericanregion(54% of theirspecies),especiallyMexico and
northern CentralAmerica.Not surprisingly, taxa adaptedto dryareas are also
relativelywellrepresented in thenorthern Venezuelan-Colombian regionandthe
drycerrado-chaco-caatinga region of the Brazilian shield.
Although representation of thesetaxa is aboutas strongin theSouthernAn-
dean regionas in theCentralAmericanregion,endemismis slightly greaterin
CentralAmerica(57% 53%vs. ofthe region's species) andeven more pronounced
in thecerradoregion(62%). Rzedowski(1962, 1978)has pointedout thatende-
mismin Mexico is moststriking amongdryarea taxa even thoughspecies of
lowlandtropicalforests dominate thecountry'sflorain termsof absolutenum-
the
bers.Despite high endemism, taxa ultimately derivedfromthesouthstrongly
predominate in theMexican dry area flora, in contrast to thenorthtemperate-
deriveddryarea floraoftheUnitedStatesdeserts(Rzedowski,1973).Such pat-
terns,especiallytheprevalenceofa preponderance ofwell-marked endemicfam-
ilieslikeFouqueriaceae,Lennoaceae,Crossosomataceae, Malesherbiaceae,and
Cactaceaeindryareas,havebeencited(e.g., Rzedowski,1962,1978)as evidence
ofa longevolutionary history ofdrytaxa,implying theuninterrupted persistence
ofdryareas at leastthrough mostoftheCenozoic. Axelrod(1979)suggeststhat
muchof theearlyevolutionand differentiation of dryarea taxa mayhave been
in edaphicallydryareaswithtaxa spreading as dryclimatesexpandedin thelate
Tertiary and Quaternary times.Whetheroriginally edaphicallyrestrictedor not,
thestrongdifferentiation ofmanyofthesegroupsin Mexicoandnorthern Central
Americaimpliesthatsome of theirancestorsmay well have arrivedvia late
Cretaceousislandhopping(cf. Bignoniaceae,tribeCrescentieaeabove, mostof
whosemembersare specializedforsuchedaphicallydrysubstrates as limestone
outcropsand seasonallyinundatedsavannahs).
It shouldbe notedthat,althoughamphitropical rangedisjunctions ofdryarea
taxa are frequent, manyof these surelyreflectrecentlong distancedispersal
(Raven, 1963),ratherthanthe ancientpatternsemphasizedabove. Moreover,
although manyofthesedryarea taxamightseemto be autochthonously Mexican
and northern CentralAmericanbased on theirpreponderance of species there,
mostof themare eitherclearlyof Gondwananaffinities or presumablyso by
phytogeographic analogy.Thus the highspecies numbersof dryarea adapted
shruband herbtaxa in Mexico and adjacentregionsare probablymostlya sec-
ondaryphenomenon resulting fromactiveevolutionary inresponse
diversification
to theincreasingly dryclimaticregimes zof thePlioceneand Pleistocene,rather
thannecessarilydue to ancientarrivalor autochthonous origin.Generadisjunct
betweenChile and California,forexample,are all primecandidatesforlong
distancedispersal(Carlquist,1982). Even some amphitropical dry-areashrubs
likeLarrea are now generally believedto resultfromrelatively recentlongdis-
tancedispersalrather thanancientdistributions (Wells& Hunziker,1976).Clearly
rangedisjunctions ofdryarea plantsmustbe interpreted on an individual basis.
Whilethesedryarea taxa are a significant and interesting componentof the
Neotropicalflora,theyare relatively unimportant in termsofoverallNeotropical
speciesrichness, just as Rzedowski(1962)notedforMexico.

MISCELLANEOUS SUBSIDIARY PATTERNS
A fewotherNeotropical phytogeographical patterns meritspecialnote.Coast-

al Brazilis noteworthy fortheconcentration of oftenprimitive speciesin a re-
strictedarea (e.g., Kubitzki,1975;Soderstrom & Calderon,1974)and some of
thearchaictaxa of coastalBrazil-e.g., primitive Dilleniaceae(Kubitzki,1975),
Bambuseae (Soderstrom& Calderon,1974),Perianthomega(intermediate be-
tweenthe two maintribesof Bignoniaceaeand perhapsclose to the ancestral
stockof theneotropical Bignonieae),a Cecropiawiththesimplyspicatefemale
inflorescence ofAfrican Musanga (Berg,pers.comm.)-maydatefromtheCre-
taceousseparationof SouthAmericaand Africa.Nevertheless no familyhas its
distributionalcenterincoastalBrazil.The samefamiliesthatarewellrepresented
thereare invariably betterrepresented eitherin Amazoniaor theAndes. How-
ever,boththerecently upliftedAndesand mostofAmazonia,whichwas under-
waterintothe Pleistocene,are relativelyrecententitiesgeologicallyspeaking,
and theapparentprevalenceofunspecializedtaxa in CoastalBrazilmaysuggest
theimportance ofthisregionas a sourceareaforotherphytogeographic regions.
Anotherratherisolatedlowlandarea notedforitsendemism(Gentry,1982a)
is theChocoregionofPacificcoastalColombiaand adjacentEcuador.Thisrich,
perhumid, butgeologically young,regionis an important subsetof whatis here
termedtheNorthern Andeanregion.The floristic significance
oftheChoco area
is almostentirely at thespecieslevelalthough itdoes have a fewendemicgenera
likeTrianaeopiper (Piperaceae)andCremosperma (Gesneriaceae).No family has
itschiefcenterofdistribution in Chocootherthanas partoftheNorthern Andean
region.
Finallythe GuianaRegion,and especiallythe GuayanaHighlands,are well
knownas areas ofhighendemismand muchphytogeographic interest
(Maguire,
1970).Geologically thisarea is veryold,andtheplantsofthetepuisummits have
had the potentialforverylongperiodsof evolutionin isolation.Nevertheless,
exchangebetweensummit floraandthelowlandforestflorathatascendsthetepui
slopes has apparently been muchmoreextensivethanonce thought(compare
Steyermark, 1979,and Maguire,1970).Even manyofthespeciesofthesummits
are sharedwiththelowlands,whichare in turnno morethana northern phyto-
geographicsubsetof Amazonia.To be sure,thereare a few strikingly distinct
endemicspecies and generain the regionthatmightbe recognizedas distinct
families-Saccifoliaceae(close to Gentianaceae),and Tepuianthaceae(close to
Rutaceae).Moreintriguing are severalnon-endemic familiesthatarefoundinthe
Neotropicsonlyin theGuayanaregion.These includeSarraceniaceae(disjunct
fromNorthAmerica), Tetrameristaceae (twomonotypic genera,theotherinAsia),
andDipterocarpaceae (see Maguire& Ashton,1978;theopposingviewthatPak-
araimeais closertoTiliaceae(Kostermans,1978)is based on weakevidenceand
is phytogeographicallyirrelevant sincetheSouthAmericantaxonclearlybelongs
to theDipterocarpaceae ancestralplexus,no matterwherethetaxonomiclimits
are drawn).Such patternssuggestancientsurvivals,notactiveevolutionary di-
versification.
A veryfewsmallfamiliesdo have theircentersofNeotropicaldiversification
in theGuayanaarea. The onlygenerally acceptedfamiliesthatseemto showthis

patternare Burmanniaceae, Podostemaceae,Triuridaceae,Mayacaceae, Thur-

niaceae, and Rapateaceae.Togethertheyaccountfora totalof not morethan
some 300 species. All are specializedforunusuallife styles,as saprophytes,
aquatics,or semiaquatics.In generaltheoverallfloristic
significance
oftheGua-
yana area, and especiallythe GuayanaHighlands,now seems verymuchless
thanearlierbelieved.Ifthe3% ofmydatasetofmonographed speciesthatoccur
in theGuayanaHighlandsis anyindication, thetotalfloraofthatregionis min-
usculeindeed,evenwhenallowanceis madefortherelatively smallarea ofupland
Guayana.On theotherhand,thefamedhighendemism oftheGuayanahighlands,
althoughmuchless thanthe 90Wosuggestedby Maguire(1970), is somewhat
supportedby mydata set. The 77% endemicspeciesof theGuayanaHighlands
is slightlyhigherthanthe similarfigureforany otherphytogeographic region
(Table 7).
Two otherregionsthatare surprisinglydepauperateinplantspeciesas judged
fromthis data set are the West Indies and the northern Venezuela/Colombia
region,withrespectively 9% and 8% of thetotalof monographed species. The
59% endemismvalue fortheWestIndiesis almostidenticalto the60%ooverall
specificendemism ofCentralAmerica,butthe24% figure fornorthern Venezuela/
Colombiais by farthe lowest such figureforany of thesephytogeographical
regions.In thislightit is clearhow Steyermark(1979)was able to achievesuch
a finescale in delimitingcentersofendemism in Venezuela,somecentersbased
on as fewas two species. Withsuch low totalendemism,even a fewendemic
speciesbecomenoteworthy.
FLORISTIC SUMMARY
To summarize Neotropicalfloristic
patterns,we havetwomajordichotomies.
The firstis betweenthe basicallyLaurasianmontanefloraand basicallyGon-
dwananlowlandflora.The second,withinthepredominant lattergroup,is be-
tweena largegroupof familieswithAmazoniandistributional centers,and a
secondimportant groupthathas distributionalcentersin theAndes,especially
thenorthern Andes,and tendsto be poorlyrepresented in Amazonia.It is now
clear thatthe Laurasian/Gondwanan dichotomyresultsfromthe separationof
Northand SouthAmericathrough mostof Cenozoic time.The fundamental di-
chotomy betweenAmazonian-and Northern Andean-centered families,although
equallyclearcut,has notpreviously beengenerally recognized.
Virtuallyall lianas and canopytreesof the lowlandNeotropicsbelongto
Amazonian-centered taxa. Canopy trees of montaneforestscome fromboth
Laurasianand Gondwanangroupswitha gradationin therelativeimportance of
thetwofromnorthto south.Epiphytes, understoryshrubs,andpalmettos mostly
belongto Gondwanangroupswithnorthern Andeandistributional centers.Many
Neotropicalherbsare fromwidespreadpredominantly northtemperate groups;
likeGondwana-derived herbtaxa theyhave extra-Amazonian distributional
pat-
terns.Vines (as opposedto lianas)are representedin bothLaurasianand Gond-
wanangroups;however,it is noteworthy thattheonlyrepresentatives ofclearly
Laurasianfamiliesto extensively invadethelowlandNeotropicsare vines(Aris-
tolochia,Cissus,Gouania) mainlyoccurring in forestedgesituations ratherthan

matureforest.Shrubsare best representedin dryareas and typicallyshow an

amphitropicalpattern centersbothin northern
withdistributional CentralAmer-
ica and thesouthernAndeanregion.
EVOLUTIONARY IMPLICATIONS
These verydistinctive phytogeographic patternssuggestradicallydifferent

speciationpatternsin different regionsand amongthedifferent adaptivetypes.
Especiallynoteworthy are thedifferences (1) betweenupland,largelyLaurasian,
taxa and lowland,predominantly Gondwanan,groupsand (2) withinthelatter
groupbetweenthepredominant Amazonian-centered andAndean-centered groups.
One striking differential is in pollinationecology.Almosthalfof thefamilies
of Laurasian-derived trees are wind-pollinated whereasvirtuallynone of the
Gondwananangiosperm families is. Manyofthenorthern Andean-centered groups
are largely,or evenpredominantly, hummingbird-pollinated (e.g., Acanthaceae,
Bromeliaceae,Campanulaceae-Lobelioideae, Ericaceae, Gesneriaceae,Loran-
thaceae-Loranthoideae, Marcgraviaceae,Musaceae, Tropaeolaceae,Zingibera-
ceae) whereasthispollination systemis ratheruncommon in othergroups(e.g.,
a few Gentianaceae,Labiatae, Lythraceae,Polemoniaceae,Scrophulariaceae
amongLaurasiantaxa; some miscellaneousspecies of Apocynaceae,Bignoni-
aceae, Combretaceae,Convolvulaceae,and LeguminosaeamongAmazonian-
centeredtaxa). Plants withratherconspicuous,oftenlarge, tubularflowers
pollinatedby specializedlargeand medium-sized bees mostlybelongto Ama-
zonian-centered families(Apocynaceae,Bignoniaceae(Gentry,1974a, 1974b),
Cochlospermaceae (Frankie& Baker,1974),Lecythidaceae(Prance,1976;Mori
et al., 1978),manyLeguminosae(Frankie& Baker, 1974;Frankieet al., 1982)),
butare also foundamongothergroups(e.g., Marantaceae,Orchidaceae,Passi-
floraceae,Zingiberaceae). Even someLaurasiangroupsthatare mostlylarge-bee
in
pollinated thetemperate zone tendto have Neotropicalrepresentatives with
otherpollinationsystems(e.g., Scrophulariaceae, mostof whose Neotropical
generahave tinyinconspicuous flowers).Taxa havingmostlyspecieswithsmall
generalist-pollinated flowers (sensu Frankieet al., 1974,1983;Gentry,1982b)are
wellrepresented in all phytogeographical groupsand showno obvioustrends.
A relateddifference betweenphytogeographical/habit groupsliesintheirprob-
able modesof speciation.WoodyLaurasiantaxa, largelywind-pollinated, have
speciatedverylittlein SouthAmerica,even in the Andes wheretheyare eco-
logicallydominant.Even in CentralAmericatheseplantshave producedrela-
tivelyfew species. Ecotypicdifferentiation is frequentin some groups,forex-
ampleamongoaks in Costa Rica wherediff&rent sets of relatedspeciestendto
be restrictedto different Holdridgelifezones (Burger,1977,pers.comm.).The
generalpatternof littlespeciationin thesetaxa is consistent withexpectations
based on thelonggeneration timesofwoodyplants,coupledwiththerelatively
recentarrivalof mostof the taxa in SouthAmericaand even southernCentral
America.It is also consistent withmodelsthatemphasizetheimportance ofplant-
pollinatorinteractions in promoting speciation,a potentialpatentlyunavailable
to wind-pollinated species.
The woodyAmazonian-centered canopytreesand lianasare thegroupsthat
showbiogeographic patternsconsistent withclassicalzoologicalmodelsof allo-

19821 GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 587
patricspeciation.Thesearethetaxathathavethekindsofdistributions thathave
beeninterpreted as resultingfromspeciationand/or survivalinPleistoceneforest
refugia.For example,Prance(1973)selectedentirely woodyAmazonian-centered
taxa (Caryocaraceae,Lecythidaceae,Chrysobalanaceae, and Dichapetalaceae)
to demonstrate thekindsof correlatedpatternsof restricted distributions-allo-
patricamongcloselyrelatedspeciesbutreplicatedin unrelatedgroups-thatfit
thepredictions ofthePleistocenerefugemodeloftropicalforestspeciation.It is
not surprising thatForero(1976) and Lleras (1978),bothworkingwithtaxa of
woodylianas,notedsimilarpatternsin theirgroups.In thesetaxa outcrossing is
therule(Bawa, 1974),chromosomenumbersare frequently high("palaeopoly-
ploidy")and oftenstablein a genusorfamily(Goldblatt& Gentry,1980;Ehren-
dorfer,1970),and hybridization is rareor non-existent (Ehrendorfer, 1970;Ash-
ton, 1969). Major shiftsin mode of pollinationmay be rare while speciation
leadingto specialization in suchmarginal habitatsas whitesand "campinarana"
or
forests seasonally inundated "varzea" or "tahuampa"forestsis a majorevo-
lutionary theme(Gentry, 1980b, 1982d, 1982e).In general,closelyrelatedspecies
show allopatricdistributions and speciationseems somehoworderlywithrela-
tivelyfewspeciespergenus(14 spp./genus on the averagewithonly19 genera
havingover 100species)and community diversity perhapsapproaching a regu-
latedecologicalequilibrium (Gentry,1982b).
The epiphytes, understory shrubs,and palmettosthatmakeup mostof the
northern Andean-centered taxa are characterized by whatappearsto have been
explosivespeciationand adaptiveradiation,almostcertainly muchofit sympat-
ric.Generaare typically large(20 speciespergenuson theaverage;at least 120
generawithover 100species).Whileit is conceivablethatmicrogeographic spe-
ciationcouldexplainmuchofthehighdiversity oftypicalAndean-centered taxa,
my (Gentry,1982a) attemptto fitthe Choco florato the expectationsof the
Pleistocenerefugemodelweredistinctly equivocalwithverymanylocal endemics
occurringscatteredthroughout the regionand constituting veritable"species
swarms"in largeevolutionarily plasticgeneralikeAnthurium, Piper,and Cav-
endishia.The samepatterns are documented by monographs ofspecificAndean-
centeredtaxa(e.g., Harling,1958;Smith& Downs, 1974,1977,1979;Berry,1980;
Luteyn,1983).Berry(1980andinprep.)suggested fromhisanalysisofspeciation
patterns in Fuchsia thatspeciationin thisgroupmightreflect"shifting balance"
phenomena(Wright,1977;Templeton,1980)withmajorgeneticreorganizations
or genetictransilience (Templeton,1980)optimizedbothby the smalllocalized
populationsand theneed forconstantrecolonization of a habitatpartitioned by
mountains, local rainshadows and otherclimaticeffects, verticallyshiftingcycli-
callycoalescingandseparating vegetationalzones,andfrequent landslides,which
regularly provideopen areas forcolonization.The relativelyshortgeneration
timesof theseherbaceousor shrubbygroups,as well as theirtypicallyrather
specificpollination relationships, shouldprovideideal conditions forrapidevo-
lutionarydifferentiation, even understableclimaticconditions.Thereis some
evidencethatspeciationin thesegroupsfrequently involvessuchphenomenaas
polyploidy (e.g., Psychotria, Hamilton,pers.comm.),hybridization (e.g., Fuch-
sia, Berry,1980)or cleistogamy (e.g., Marcgraviaceae,Bedell, pers. comm.),
Microgeographic, perhapseven moreor less sympatric, speciationis probably

the rule ratherthanthe exception.Shiftsin specificpollinators are a common

modeof speciationand co-evolution givingrise to finerbiotictuningofprecise
plant-pollinator systemssuch as thosein Heliconia (Stiles, 1975),Orchidaceae
(Dodson, 1975),Ericaceae(Luteyn,pers.comm.)orFuchsia(Berry,1980)seems
a majorevolutionary theme.In suchgroupsspeciationwouldappearan altogether
open-ended phenomenon without theslightest hintofanykindofecologicalequi-
libriumor limitson speciesdiversity.
The Neotropicsaremuchricherinepiphytes thanthePalaeotropics (Richards,
1973;Madison,1977;Burger, 1980).However, the tendency has been to explain
thisdifference as due to lack of extinction in the relatively(at least to Africa)
constantly mesicNeotropicsand to interpret highepiphyte diversity as reflecting
a longhistoryof mesicconditions(Burger,1980). proposethathighdiversity
I
in epiphytesand otherNorthernAndean-centered groupsresultsmostlyfrom
recentverydynamicspeciation,almostthe antithesisof theprevalentlack-of-
extinction hypothesis.Ratherthanthefloraof tropicalAfrica(and to a lesser
extentAsia) beingimpoverished withrespectto theNeotropics,thelattermay
be considered as uniquely and phenomenally enriched.
Thereis increasing circumstantial evidencethatthiskindof unusuallyrapid
speciation,concentrated alongthebase and lowerslopesofthenorthern Andes,
has involvedmuchco-evolutionary interaction and has not been restricted to
plants.Thus Stiles(1981)has shownthatthereare over400 species of flower-
feedingbirdsin theNeotropics,including 315 speciesofhummingbirds alone,as
comparedto 100-150flower-feeding speciesin each Palaeotropical realm.More-
over,the Neotropicalflower-feeding birdsare generallymuchmorespecialized
and show muchgreaterflowerspecificity thantheirPalaeotropicalequivalents.
Stiles(1981)suggested fromthesepatterns thatbird-flower coevolutionprobably
beganrelatively earlierin theNew Worldthanelsewhere.However,in thecon-
textofthebotanicalpatterns discussedinthispaper,I wouldsuggestinsteadthat
it is probablethatbird-flower coevolution,in general,and hummingbird specia-
tion,in particular, has been muchmorerapidin theNeotropics.Certainly hum-
mingbirds are concentrated in tropicaland premontane partsof the northern
Andeanregion(134 species in Colombia,'ca. 133 in Ecuador (Bleiweiss,pers.
comm.),97 in Venezuela,118 in Peru (Parkeret al., 1982),comparedto 52 in
Panama(Ridgely,1978),51 in Costa Rica, 37 in Guatemala(Land, 1970),60 in
Mexico; see Stiles,1981,table5, forecologicaland altitudinal distributional pat-
terns)exactlyas aretheAndean-centered, largelyhummingbird-pollinated groups
ofplants.
Althoughthedata forflower-visiting bats are less precise,therathergener-
alized distributional patternsshownby Koopman(1981),are clearlysimilarto
thoseof flower-visiting birdsand myAndean-centered planttaxa. The greatest
concentrations ofnectar-feeding bat species(13 specieseach) are in thenorthern
Andeanregionand CentralAmerica.
It is notyetclearwhethersuchpatternsare morea cause or an effectofthe
apparentevolutionary explosionofplanttaxa characteristic ofthenorthern An-
dean region.However,Terborghand Winters(1982) have shownthatforbirds
in general,local endemism is strikinglyconcentrated on thewesternside of the
northern Andes,andKeisteret al. (1983)have suggestedtheoretical reasonswhy

thepartitioned populationstructures prevalentin thisgeographicalregionshould

favorrapidcoevolution.Moreover,Marshallet al. (1982)notea grosslysimilar
patternformammalswithformation oftheIsthmianlandbridgeresulting inwhat
theyconsiderto have been a balancedexchangeof equilibrium faunasbetween
Northand SouthAmericafollowedby a uniqueand unbalancedsecondarydi-
versificationof theimmigrant taxa in SouthAmerica,apparently somehowas-
sociatedwiththeAndes. Perhapsthe acceleratedratesof speciationhere sug-
gestedfornorthwestern SouthAmericanplantsis partof a muchmoregeneral
phenomenon.
To summarize, theexceedingly dynamic, evenexplosiveevolutionthatseems
to characterizethenorthern-Andean-centered taxa has givenriseto a verysig-
proportion
nificant ofthetotalNeotropicalflora.Almosthalfof all Neotropical
plantspecies appearto belongto thesegroupsof epiphytes, understory shrubs,
andpalmettos, all ofwhichare muchmorepoorlyrepresented in thePalaeotrop-
ics. Thus thehistoricalaccidentof theAndeanuplift,withtheconcomitant op-
portunityforexplosivespeciationamongcertaintaxa ofGondwananplantshav-
ingtheevolutionary potentialforexploitingepiphytic,palmetto,and understory
shrubstrategies, maylargelyexplainthe"excess" plantspeciesdiversity ofthe
thisapproximately
Neotropics.It is essentially halfoftheNeotropicalflorathat
is missingin thePalaeotropics, althoughsimilarpatternson a smallerscale might
be expectedin New Guinea,whichseemstheclosestPalaeotropicalequivalent
oftheAndeancordilleras.
CONCLUSION
A richangiosperm florasimilarto thatintherestofthetropicsevolvedduring

thelast halfoftheCretaceousin SouthAmericabutthisflorahas subsequently
givenriseto manymorespeciesin theNeotropics.
At theend of theCretaceoustherewas a possibility forrelatively directflo-
risticinterchangebetweenSouthAmericaand tropicalNorthAmericavia island
hoppingalong the proto-Antilles; manyof the Neotropicalgroups,especially
some of thedryarea taxa thatshowstrongdifferentiation in bothregions,may
reflectthisearlyinterchange.
Upliftof the Andes,mostlyin Neogenetime,led to an incredibleburstof
speciationin a numberofGondwananfamilies.A similarevolutionary explosion
inthesametaxaalso tookplace inCosta Rica and Panama. The taxonomic groups
thathave undergonethisevolutionary explosionhave distributional centersin
the northern Andeanregionand southernCentralAmerica,are poorlyrepre-
sented Amazonia,and consistmostlyof epiphytes,shrubs,and palmettos;
in
theirpollinationsystemssuggestthatcoevolutionary relationships withhum-
mingbirds, bats,
nectar-feeding and perhaps such specialized bees as Euglossines,
have played a prominent role in theirevolution.The evolutionary phenomena
associatedwiththeAndeanupliftaccountforalmosthalfofthetotalNeotropical
floraand are thuslargelyresponsible fortheexcess floristic richnessoftheNeo-
tropics.
Closingof thePanamanianisthmusin thePlioceneled to (1) southwardmi-
grationof someLaurasiantaxa intotheAndeswheretheyhave becomeecolog-
icallydominantdespiteundergoing littlespeciation,at leastin woodytaxa, and

invasionof lowlandGondwanantaxa of canopytreesand lianas

(2) northward
intoCentralAmerica,leadingto theirecologicaldominancein lowlandtropical
foreststhroughout speciationinCentralAmer-
theregion,despitelittlesignificant
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Neotropical Floristic Diversity A, Gentry

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Neotropical Floristic Diversity: Phytogeographical Connections Between Central and South

America, Pleistocene Climatic Fluctuations, or an Accident of the Andean Orogeny?

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The interchangeofplantspeciesbetweenNorthand SouthAmericahas been

Severalmajorgeologicaleventshave had profoundeffectson the evolution

1 Thispaperwas presented inbothSymposiapublished inthisissueoftheAnnalsoftheMissouri

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ical Americanangiosperm familiesas Bromeliaceae,Humiriaceae,Cactaceae,

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pinchingoffof a segmentof thePacificplateas the Caribbeanplate,withrela-

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representshareddescentfromhypothetical commonwide-ranging middleCre-

thispaperin a broadsenseto includeMexico.

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mentedby van der Hammen(1974) and his associatesfromthe palynological

One oftheoutstanding featuresoftheNeotropicalflorais itsextremerichness

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Whileit is generallyappreciatedthatAfricais floristically depau-

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examplecomesfromtheRio Palenquefieldstationin westernEcuadorwhere,

The greaterdiversityin the Neotropicsresultsin partfroma conspicuously

5The discoveryofa primitive in SouthAmerica(Maguire& Ashton,1977)does not

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species each. Unlike most of theirPalaeotropical counterparts,manyof the Neo-

The phytogeographical significanceofthenewdevelopments inplatetectonics

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tAceraceae 1/5 (Krameriaceae) 1/15

otherpredominantly Laurasianfamilieswere suggestedas probableearlierar-

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No. Spp. No. Ind. No. Spp. No. Ind.

sian?) genus, and the familymay be fundamentallyLaurasian in origin.

and Gondwanan elementshas not greatlyincreased overall Neotropical floristic

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TABLE 4. Amazonian-centereaGondwanan families.NumbersindicateNeotropicalgenera with

Anacardiaceae (?) 15/133 (+2/257) (Lacistemaceae) 2/14

elements.Similarlyan Oligocenesite in PuertoRico (Graham& Jarzen,1969)

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TABLE 5. Andean-centeredGondwanaland groups. Numbers indicate neotropicalgenera with

Total 199/4,218 (+10/1,800)

t = does not reach CentralAmerica.

ness of the lowlandtropicalrainforestdecreasesdramaticallynorthward. My

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TABLE 6. Miscellaneous taxa. Numbersindicateneotropical

Guayana-centeredgroups Ceratophyllaceae 1/2

finedsouthward"bytheslightly lowertemperatures duringglacialadvances(Ax-

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TROPICAL AMERICA Ldyer 1ft Md) Fkxd NewIr opct ki

FIGURE1. TypicalAmazon-centered distributionof a taxonof canopytrees,Moraceaetribe

withthechanging altitudinalzonationofthevegetation broughtaboutby climatic

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FIGURE 4. Neotropical regions.

weedyand theirpatterns are notso wellmarked.Nevertheless, exceptfora few

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The Amazonian-centered taxa include38%oof the totaldata set of mono-

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not even Dilleniaceae,whose strongrepresentation in the coastal regionwas