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 Reviewed work(s): Source: Annals of the Missouri Botanical Garden, Vol. 69, No. 3 (1982), pp. 557-593 Published by: Missouri Botanical Garden Press Stable URL: . Accessed: 05/02/2012 13:01
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The interchange ofplantspeciesbetween North and SouthAmerica has been a majorfactorin determining the Neotropical obphytogeographical patterns servedtoday.Although thishas longbeen realized,recent historical geological evidenceas wellas increasing knowledge of today'sdistributional patterns now makespossiblea morethorough ofhow thesepatterns analysis This originated. reviews thegeological paperbriefly summarizes thecomposition background, of outthestriking ofmany theextant ecological Neotropical flora, points consistency taxa and lifeforms, and suggests how some of the present phytogeographical have developed from patterns theinterplay ofthesefactors.

Severalmajorgeological eventshave had profound effects on the evolution and distribution oftheLatinAmerican flora. One ofthemostsignificant ofthese was the separation of SouthAmericafrom Africathatbegan only 127 million yearsBP (Rabinowitz, 1976;McKenna,1981)with contact or near-contact presentuntil 80-90million yearsBP (Raven& Axelrod,1974;McKenna,1981).Duringmostofthefirst third ofangiosperm evolution, a timeduring whichmany of themodern orders andfamilies ofplants arose,SouthAmerica was a partofthe WestGondwanaland cradleoftheangiosperms (Raven& Axelrod,1974).However,during all oftheTertiary virtually andmuchoftheCretaceous SouthAmerica was an islandcontinent, at leastfrom theperspective oftropical plants, and mostoftheevolution ofitsrichand varied flora tookplace in isolation following separation from Africa. Thus,suchcharacteristic andecologically important trop1 Thispaper inbothSymposia was presented inthis published issueoftheAnnals oftheMissouri Botanical Garden-theSymposium on PlantGeographical of Changing Results at CenozoicBarriers theXIII International in Sydney, Botanical Congress Australia, 1981;andthe1981 Systematics Symof theMissouri posium Botanical Garden.As used herein, references documented as "thissymposium"include papersfrom bothsymposia. 2 Thispaperstems from largely insights developed during field work overa number ofyearsin various partsofLatinAmerica, and supported by theNational ScienceFoundation (GB 40103, INT7920783, DEB-8006253, DEB-UT-20325, DEB-8006253), theNational Geographic Society, andUSAID I thank D. Axelrod, (DAN-5542-G-SS-1086-00). P. Ashton, P. Raven,J. C. Dodson,A. Graham, Rzedowski, J. Terborgh, and B. Simpson forreviewing the manuscript and numerous colleagues including D. 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. C. Stace,P. Taylor, Soejarto, W. Wagner, D. Wasshausen, H. Wilson, andD. Wunderlin for providingdataon Neotropical speciesnumbers or distribution patterns oftheir taxonomic specialties. 3 Missouri Botanical P.O. Box 299,St. Louis, Missouri 63166. Garden,

ANN. MISSOURI BOT. GARD. 69: 557-593. 1982.

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[VOL. 69

ical American angiosperm families as Bromeliaceae, Humiriaceae, Cactaceae, and Caryocaraceae, had ampletimeto evolveand radiate in SouthAmerica. Although thehistory oftheseparation ofSouthAmerica from Africa is clear, the tectonics of the earlierseparation of NorthAmericafrom the Gondwanan landmassis still uncertain (see Coney, thissymposium). Apparently North America (including the CentralAmericanpeninsula)separatedfromAfrica-South in Jurassic America time, wellbefore theorigin oftheangiosperms (Lillegraven et al., 1979).However,by late Cretaceous a chainof volcanicislands(Nicoya haddeveloped Complex) connecting Central America with Ecuador(Dengo,1975; Lillegraven et al., 1979).New geologicalevidence(Dickinson & Coney, 1980) suggests thata connection between nuclear Central America and SouthAmerica was reestablished inLate Cretaceous as bothmajor American landmassesmoved westward moreor less in tandem. How muchof the late Cretaceousconnection betweenCentraland South America was above sea levelis notknown, although itprobably consisted mostly of an interrupted islandarc. Further the picture, complicating muchof northwestern SouthAmerica was submerged during mostof the Cretaceous(Irving, 1975).Boththelow coastalrangeofwestern Colombiaand southern Darienand thenorthern AndeanCordillera Occidental wereoriginally islandarcs associated withwestward movement of the SouthAmerican plate during upperMesozoic time(Zeil, 1979: 193). McKenna (1981) has suggested thatthe present Pacific coast of northern SouthAmericamayhave resulted from accretion the during earlyTertiary ofleft-behind fragments ofthesouthend oftheCentral American volcanicarc. According to McKenna(1981) some oftheislandsofthisarc may have remained above water,separated from NorthAmerica onlyby sequential and closing of watergaps,untilcolliding opening withSouthAmerica and proa plausible viding scenario for"Noah's Arc" dispersal ofhystricognath rodents betweenNorthand SouthAmerica, a modelconsistent withthe discovery by Juteau et al. (1977) thatmuchof the west coast of Ecuador is formed from a blockof originally oceaniccrust. Therewas a general regression ofepicontinental seas from northwestern South America at theendoftheCretaceous (Harrington, et al., 1979) 1962;Lillegraven concomitant witha late Cretaceous thatgave riseto theforerunner orogeny of theColombian Cordillera Occidental (Irving, 1975).Morethan160kmwas added to thenorthwestern SouthAmerican continental Late Cretaceous margin during time.The Greater thenfarto thesouthoftheir Antilles, and in present position part also constituted submerged, oflateCretaceous presumably partofsomekind inter-American connection as suggested by'Malfaitand Dinkelman (1972) and discussedin the context of biogeography by Tedford (1974) and Rosen (1974). a different (See Pregill (1981)for butmuchofthegeology on which interpretation it is based (Perfit & Heezen, 1978)has apparently been superseded (Dickinson & Coney, 1980;Coney,thissymposium).) to thisinterpretation, the According continued westward movement ofSouthAmerica andtheslightly southwestward movement of North America led to decoupling faults thatseparated theprotoAntilles from bothland masses. A new subduction zone thenformed to their northeast as theproto-Antilles movednortheastward. Continued westward movement of SouthAmerica and southwestward movement ofNorth America led to

and northern southern withboth strong of the West Indianflora.thepresenceof a previous. from to the of now locally intheearly presence Tertiary judge beenevenstronger there taxa like Aetanthus (although American extinct otherwise exclusively South in from the northern this generalized be separating identification problems may in Europe in the Eocene.thissymposium). south moredirectly an original position platefrom withtheCaribbean eastward North (Dickinson and SouthAmerica partofthegap between ofMexico. and Nyssa. the earlyCenozoic. composition of a protostocking would be expectedas the resultof an original affinities.closing and trench American oftheCentral formation & Coney. biota. Engelelements 1969)alongwithnorthern ham & Jarzen.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 559 plate. In thelateTeritary. mostly presumably continents. inter-American formoreor less direct taceouswouldhave had the opportunity of Panama. North andSouthAmerica ofPanamabetween oftheIsthmus closing Although theLatin for Cenozoicbarriers" "changing one ofthemost important was clearly late Cretaceous albeitinterrupted.although Thisdichotomy mayhave evidence with thegeological (Coney. to takeadvantage ofisland and in a position in WestGondwanaland wereextant America. flora American likethemodern tropical seemsto havebeenmore of the apparent affinities to judge to whatextent However..may at varioustimesduring tively long distancedispersal betweenthe two insteadmoreor less directlate Cretaceousmigration reflect the dichotomous via islandhopping.withrelaplateas the Caribbean of thePacific offof a segment pinching to South of the whole Caribbeanregionrelative displacement tivelyeastward also moved fault south oftheMotagua America Central Probably nuclear America. was completely linkbetween North and SouthAmerica Thattheinterrupted northbegantheir as theproto-Antilles oftheTertiary. and nuclearCentral betweenSouthAmerica land connection Americawithsubstantial coalesced intotoday'slowerCentral onlyabout in thePlioceneapparently ofPanamaestablished acrosstheIsthmus et al. America. Southand North between stepping-stones American families or predominantly Thus mostof the cases of exclusively inbothtropical and North differentiated components strongly likeCactaceaewith forby chancerelaRaven and Axelrod (1974) accounted which SouthAmerica. American thecontinents fairly shorter water gapsbetween wouldhaveprovided connection of importance our conceptofthephytogeographical earlyand thusmaymodify ofplants time families andgenera modern In lateCretaceous many this lateevent. 1982). 3 million 1978. by locatedbetweenNorthand SouthAmericaas emphasized Antillean region do notaccord land connections his arguments fordirect Rosen (1974). Similarly.Marshall yearsago (Keigwin. acLoranthus pollentypethatwas alreadypresent in ofPuerto Rico (Grathe Oligocene and Catostemma to cording Muller.1980). than itis today. ofislandsin theregion againled to uplift an associatednewepochofvolcanism These islandseventually is verydifficult floras might and SouthAmerican Paleoceneand earlyEocene NorthAmerican . whichhave neverbeen able to reachSouth hardtia. In of the Isthmus priorto the Plioceneformation islandhopping North America ofwhatis now temperate theflora oftheearlyTertiary general. Hauya. at thebeginning ruptured in theCreextant meansthatonlythoseveryold taxa already wardmovement. Liquidambar.1981) likeFagus.

and a finalshift of the highlands the live oak forests 1972).or whether taceousearlyangiosperm betweenSouth Americaand tropicalNorth interchange tivelydirectfloristic The difficulty and successive in potentially Paleocene flora They findthe generalized the earlyTertiary. interchange centers suggestedfor such genera as Ocotea. othersalso occurring noullia.perhapsvia islandhopping remained of SouthAmericaand NorthAmerica floras the-Tertiary connection.and Swartziaby Leopold and MacGinitie withthepossibleexAmerica thanin Central todayin SouthAmerica resented and two America. Luehea. Dilcher.1974. as endemic and suggested fossil theTertiary in SouthAmerica. Beilschmiedia. thispaperin a broadsenseto include is usedthroughout 4Central America .560 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL.1974). Central to thoseofthesubhumid floras to of northern Mexico. migration inter-American direct via a relatively distributions present to theEocene. modern pan-neotropical suchgeneraachievedtheir of establishment it was longbefore that evidencesuggests and ecological graphic or mamlandbridge. theysuggest stocks. there). work (cf. ofAfrica totheseparation subsequent America in identaxonomic errors is compounded by theserious offloristic interpretation muchearly thatare now knownto have characterized of fossilfloras tification 1975).Unfortunately well or better represented to show when Americaand SouthAmericais inadequate Central recordfrom butphytogeodistributions.thefloristic by Leopold and MacGinitie As summarized America North tripartite biotically theonlypartofthen region.whilethe in tropicalAmerican and autochthonously floraanalyzedas characteristically flora American North an olduniquely either tropical havetorepresent would origin to accountfortheoverwhelm(whichlatermusthave spreadto SouthAmerica today)or earlyfloristic ofmostofthosegenera distributions ingly pan-American distributional American4 totheCentral Contrary with SouthAmerica. sumably prior reachedCentral floristic elements evidently some SouthAmerican Although alongthe late CretaceousAntillean Americaearly. paleobotanical of the affinities (1972).Graham. rep(1972). mesophytic mixed and warm-temperate tropical to flora theOligocene American tropics. Mexico. Cedrela. changesthrough that but suggest region. forests conifer Cordilleran modern the latest most of the generasharedwithSouth Americaduring Presumably ancescommon Cretaceous reflect wide-ranging and earliest Tertiary Cretaceous Eocene late middle and subhumid tors(cf. 69 middleCrewide-ranging common hypothetical shareddescentfrom represent relaan independent.Beilschmiedia. intheMiocene(Leopold& MacGinitie. RockyMountain directcontactwithnuclearCentralAmerica. phytogeographic to relateto thatof any extant difficult Asian subrelatedto the southeast was primarily themesic earlyEocene flora andlateEocene themiddle forest. Manyoftheseplantshave wind-dispersed thePanamanian forlongdistancediscandidates and are unlikely seeds or fruits mal-dispersed withhavingachievedtheir 1982a). of theeight speciesonlyin Central of Cedrela(three ception Berfour three of the Even genera-Homalium.and Engelhardtia-related to Rocky Mountainfossilforms are as in their American America to Central ranges. preroute.Hickey& Wolfe. Raven & Axelrod.whichwouldbe consistent persal(Gentry.all are better Oreopanax. 1972:8.

as elegantly docuthree of the temperateNorth American genera-Alnus. 1968). picture of majoruplift of the alreadyextant southern and central Andes in the and ofthenorthern Mid-Cenozoic Andesmorerecently seemswell established. 1974. Further southin Central in thenrecently America. theopportunity forlimited late Cretaceous migration between Southand North America wouldreadily accountforthemanycharacteristically Gondwanan taxa withdistinctive and strongly differentiated CentralAmerican derivatives (see below). similar to thedeciduous forest ofthe eastern UnitedStatesand todayrestricted to intermediate elevations. Pollenof ten of the fourteen arborescent generathatwouldhave been expectedin similar depositsin the southeastern UnitedStatesis present. the Paleocene pollenflora of Colombiacontained suchlowlandtropical exclusively elementsas Annonaceae. none of theSouthAmerican taxa thatbarelyenter Central America are represented in theWestIndieseither. In themontane tropics these -climatic oscillations tooktheform of an altitudinal and compression lowering of vegetational zones. As might be expected.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 561 fundamentally different (Germeraad et al. ofthenorthern Andestookplace onlyin thelastfive million or so years.Although theAndeanorogmorecomplicated enywas certainly thanoften thegeneral supposed(Zeil. The secondCenozoicgeological eventwithmajorphytogeographical importancefortheNeotropics was theuplift oftheAndes. whilea basically North American plant community.The manywidespread SouthAmerican families and generathattodaybarelyenter Central America in eastern Panamaprovide evidence that theTertiary barrier to northward migration was equallyeffective.).The Andesare unique:by farthemostextensive mountain rangein theworld'stropics. Whileredeposition ofpollenfrom at higher plantsgrowing altitudes couldhave affected theseresults (Axelrod. is clearthattheeastern deciduous forest elements wereat least present in theregion. 1973). was well represented. pers. North gapbetween and SouthAmerica twovery separated distinct Cenozoicfloras inthetwocontinents despite a probable early opportunity forrelatively direct islandhopping across theproto-Antillean chain. emerged Panama. and several generaofpalms:Mauritia.Bombacaceae. The Colombian Cordillera Central is olderand existed in theCretaceous already (Zeil. (Hammen & Garcia. 1982) analysisof the Paraje Solo formation suggests thatin the Miocene the southern Veracruz lowlands had onlya fewscattered representatives oftheSouth American-derived tropical forest whichnow characterizes the region. Araceae.In contrast..1966).However. comm. and severalpollengenera Astrocaryum. A third eventof majorphytogeographic importance was theadventofPleistoceneclimatic fluctuations associated with glacialadvancesandretreats at higher latitudes.These samethree generaappearin theSouthAmerican palynological record onlysubsequent to closing oftheIsthmianconnection in latestPlioceneand Pleistocene times. 1979: 109)butprobably was erodeddownto a low rangeof hillsprior to therecent Most of theuplift orogeny.only are present in the Miocene Gatunformation and none of themare present in earlier Panamanian deposits (Graham.1979).For exampleGraham's(1973.during Plioceneand Pleistocene times (Hammen. 1979). To thewater summarize. and Myrica- .Flenley.

1973.have long-lived Pleistocene in therelatively time short and lianashad adequatetime to speciateso profusely Moreover of thePleistocene? and fewgenerations availablesincethebeginning forformer refugiathathave been citedas evidence patterns thedistributional contact zonestaxacoming at coincident together unrelated speciespairsinmany in situspeciation Nevergradients. generally FLORISTIC BACKGROUND richness flora is itsextreme oftheNeotropical features One oftheoutstanding ofNeotropical fornumber plantspecies in species. Palaeotropical muchhigher Brazilaloneandthebestavailableestimate Amazonian timates 30.000(Gentry.g. perhapsslightly have generally acceptedRaven's estimates." popularconception forestspecies repeatedly for speciation. auPrance(1977) and other fortropical and 21/2 timestheestimate Australasia. Simpson. thors are availableforthePalaeofloristic diversity forcontinental estimates higher.1978). as populations of tropical conditions withthe and recoalesce(see papersin Prance.000 .1982). during dry periods riphery ofAmazonia Haffer.Nevertheless (Jacobs. trary to earlysuggestions and subtropical ofaridtemperate periods pluvial notthesameas thewellknown wet periodswere glacial advances. circulation that enough spheric forests neotropical the lowland precipitation changes in these cyclical resultof in extent aroundthepepockets.000speciesfor one is 18.000. from alongenvironmental could also result bybiogeographers modelhas beenwidely accepted thePleistocene refuge theless Not onlydid formuchtropical speciation. speciesfor 25.1978.tropical desertsthataccompanied pluvialperiods.Some problems fragment trees refuge modelhave been noted. In thelowland Cordillera record in theColombian but changesin precipichangesassociatedwithglacialadvanceswere minimal wereassociated The cyclesofglacialadvanceandretreat werepronounced. comm. 1978). as a generalexplanation accounting islands. effect multiplicative lackedsuchrefugia. Haffer..69 the palynological mented by van der Hammen(1974) and his associatesfrom temperature tropics Oriental. Simpson & symposium.)esthanfortheentire region. Other recent.1965. alone(Brennan.verydifferent of thisdynamic The biologicalsignificance it is that optimal provides ofthe stable"forest primaeval.000-30.000 is number of plantspeciesfortheNeotropics it seems clearthattheestimated Prance(pers. Africa 1979)and tropical speciesforcontinental tropics-30.this the from model.562 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL.ThusRaven's(1976)estimate Africa for times as high as hisestimate plusMadagascar tropical three was 90. chiefly to scattered reduced wereperiodically Prance. forCentral 1978)and an (overlapping) to 20.butthe survive dryperiodsin thesehabitat forest species successfully have multiplied couldpotentially fragmentation range cyclesofmultiple repeated In other theNeotropics have many speciesbecause ofthe if perhaps180?out of phase withthe temperate-subtropical atmomuch the of normal water the earth's of the glacialadvancestiedup so for As a not was left "normal" tropical rainfall.For example. tation Conrespectively. dryand wetperiods withalternating lowlands in thetropical were ofthetropics 1970). 1971. forMexico is 14.000(Rzedowski. 1969.000 America to 20. thewetperiods (e.Instead. their Africahas few because it of its more numerous refugia. (Haffer.000 theFloraMalesianaregion 1974). words.

Astragalus. and Buxaceaefrom and in the to theNeotropics usefulonlyforgenerarestricted bers are directly region.200Neotropical missed. (mostly (1980). species.'s (1976) Actually. poorNeotropical relatively with zone genera temperate Ipomoea) or arebasically an overalltotalof implying ofthetotalspeciesof thesegeneraare Neotropical. speciesover10cmdbhas evidence. Raven's by rich as suggested species plant seed all of a list Neotropical I compiled First. Gentiana).000+ speciesare either The remaining tropical. tropical genNeotropical non-endemic 533non-monographed theother Unfortunately.000Neotropical The suchan estimate. floras generabased on the availableregional for herbarium Garden Botanical Missouri theentire by a searchthrough mented were a fewsmalllocal genera Although from theregion.1973.Brennan. incompiling serious problem is another there However. (1977).1977)have suggested Asia (Whitmore. nearManausis as diverse offorest of 179treespecies. as neotropical or moretreespecies as many by Raven's estimates? morespeciesas suggested have as many really are verytentative.. 40. 7.I estimated complete.000-10. plantspecies remain of the worldand verymanyNeotropical species plant all of quarter a probably Choco. have species oftheendemic (e.orbasically genera pantropical groups-large into twomain About20.manystudying thatthedipterocarp 1975.g. are the of thatregion forests sample recent et Prance al.). Carex.Gramineae Moldenke forspeciesnumfigures myowndata.000speciesbased on theWillis erainclude likePiperorEugenia. a totalof60.15 cm dbhin a hectare have Asianforests Southeast Ifindividual Asianforests.000) a third that suggesting Neotropics in cosmopolitan genera(e. A fewadditional inmostother 1973) Heywoodet al.and Bignoniaceae.Leweretakenfrom used forsomegroups-Compositae fromPolhilland Raven (1981). of tree of number world's citing samples richest.Ashton.theresulting probably ofNeotropical speciesineach thenumber Second. (Airy by Willis'sDictionary supplied andthefigures wereavailable.660generawithalmost65. diversity floristic ofNeotropical The availableestimations as incredibly arereally theNeotropics whether accurately to assess more In order countthe to to try decided I estimates.000Neodata set accounted The resultant species. as mostoftheSoutheast Neotropics the can their equivalents.. fall Thesegenera figures.A well-documented been described (Gentry.whentreatsuchas theFloraNeotropica monographs usingrecent genus. ments were datasourceson speciesnumbers cases. OrchidaceaefromDressler mostly guminosae D'Arcy (1979). Colombian likethewestern regions likeAraceae or Ericaceaealmostnone and in manygroups endemic are strictly 1982a). Solanaceae from Davidse. sonably series. fora genusare givenby geographical fewcases wherespeciesnumbers forover3.comm. directly.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 563 depaurelatively is floristically thatAfrica appreciated Whileit is generally Southeast of flora the perate(Richards. names generic additional generaseemsrealistof over4. parts other than floristically known much more poorly are generally Neotropics In undescribed. Draba. Verbenaceaeand Eriocaulaceaefrom (1981).. at least76.1964. Shaw.pers. The Willis Sabiaceae.000of Neotropics.1979).g. species of number suppleand familial monographs. Perhaps only 10-15% representation .e. Laurasianherbsthatrangesouthintothemontane in the thattendto be wellrepresented genera thesespeciesfallintopantropical are Neoofthem to a half(i.

e.Pandanaceae (700spp. broadfamilial traditional retaining species..000 & Ashton. (e. with essentially via longdistance Neotropical dispersal) arrived recently torsprobably relatively have of thesefamilies witha totalof 5. and two-Bromeliaceaeand Cactaceae-have about 2. endemic" of "essentially bythedefinition ofthefamily thestatus change . Africa. a local florula in theprocessofwriting and recollections in a minuscule1. connections. Even of endemic complement important and moreecologically larger or there are at least38 endemic delimitations. Gentry. on different on thepartofworkers of in thecontext similarities floristic pointof thesepantropical The relevant thepresent flora Neotropical that distinguish is that thepeculiarities thisanalysis stocksharedat least withAfrica(i. ENDEMIC FAMILIES a conspicuously in partfrom in the Neotropics results diversity The greater families.mostofwhoseancesendemic one or twoAfrican (i..). genusthanlike otherAfrican betweendifferent of generain common of percentages Statistical comparisons sincetheimportant misleading areas (e.1973)are especially tropical disproportiongeneracontribute generatendto be sharedwhilesmallsegregate parochialism by taxonomic is also confused The picture atelyto thedifferences.and more than50 species-Dipterocarpaceae5 have over families Neotropical endemic (68 spp. subsequent were discovered almost total to the taxa now brings problem of tentatively identified evaluation must that there suggest figures Such Rio Palenque. Thorne.. or to 86. floristic have arisendespitea common Asia as well.. from 100 species described the that would elevate neotropical plantspecies undescribed be at least 10. ones (Hladik & Halle.564 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL.).are also found are morelikesomeNeotropical moreAfrican and many genera in theNeotropics in prep. 1973. theRio Palenquefieldstation examplecomesfrom 61 newspecies 1978). In this withtropical and to a largeextent West Gondwanaland) was commonality whether thisintercontinental contextit is almostirrelevant movements.000 seed species plant ofneotropical totalnumber of90. than generally For example. (Dodson & Gentry. 1977)does not (Maguire in SouthAmerica dipterocarp ofa primitive 5The discovery used here. was fairly anyserious beyond seemsestablished partsoftheworld plantspeciesthanother doubt.000 species estimate neotropical that Raven's original We mayconclude in richer Thattheneotropics as a wholeare extraordinarily accurate.g.). 1974). The questionto be derivedfrom dispersaleventsor continental more stockhas givenriseto so many addressed hereis whythissharedfloristic speciesin theNeotropics.690species (Table 1).30% of thegenerathatoccurat Makokou.g.).000 more. Six essentially Nepenthaceae a hundred species. Good. pantropical showsverystrong generally flora The Neotropical is wellknown families tropical among predominate distributions Thatpantropical areevenstronger similarities thesephytogeographic However.e.Gabon. 69 in western Ecuadorwhere. continents.7 km2area. with level and especially at the generic especially realized. Thirteen families families with to onlythree endemic palaeotropical over50 speciesas compared (580spp.

family The other families listed byPrance(1978)as endemic are segregates (e. 1) Tropicalforest taxa* Bromeliaceae (46 genera/2.) (1/1) Malesherbiaceae (1/27) Myzodendraceae (temp.) in tropics) Tepuianthaceae (1/5) Thurniaceae (1/3) * I havefollowed traditional familial limits. Rhabdodendraceae (2 species)was treated as a distinct in FloraNeotropica. listed(Pontederiaceae).S. erroneously or both(Heliconiaceae). Endemic Neotropical families (insomecases with a single African speciesora single monotypic or ditypic African genuspresumably recently arrived bylong-distance dispersal).000) (a widespread and Ceylon speciesalso in Africa with at least 1 derivative in Madagascar) Julianaceae (2/5) (butprobably an artificial group) Koeberliniaceae (1/1) Krameriaceae (1/15) Loasaceae (12/266 (plus 1 in Africa)) (severalsmallgenera in MexicoandSW U. in Africa)) Caricaceae(3/29 (plusone ditypic African genus)) Caryocaraceae (2/24) Cyclanthaceae (11/178) Dialypetalanthaceae (1/1) Duckeodendraceae (1/1) Humiriaceae (8/46 (plus 1 sp.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 565 TABLE 1. 5) Miscellaneous (aquatics or semiaquatics and Guayanan edaphic specialists) . 108species(plus 1 sp. Peridiscaceae). in Africa)) Lacistemmaceae in Flacourtiaceae (2/14) (included in FloraNeotropica) Marcgraviaceae (4/125) Quiinaceae (4/53) Rapateaceae(15/79 African (plus 1 monotypic genus)) Vochysiaceae (7/182 (plus 1 ditypic African genus)) Trigoniaceae/Trigonia (1/24) Bignoniaceae/Bignonieae (46/359) Lecythidaceae/Lecythidoideae (300) Brunelliaceae (1/51) Calyceraceae (4/46) Columelliaceae (1/4) Gomortegaceae (temp.) Martyniaceae (3/13) 4) Taxa of drytropicallsubtropical parts of both continents Cannaceae(1/55) Cyrillaceae (3/13) (?) (in Europein Eocene.. mostly WestIndian) Mayacaceae(1/9 (also 1 in Africa)) Saccifoliaceae (1/1) Sarraceniaceae (?) (3/17) (only1 genus(6 spp.g.) (1/11) Nolanaceae(1/18) Tovariaceae in Capparidaceae (1/2) (included in FloraofPanama) Tropaeolaceae (2/92) 2) Taxa of dryandlor Andean parts of South America (a few reachingNorthAmerica) 3) Taxa of dryparts of tropicalNorthAmerica Crossosomataceae (1/4) Fouqueriaceae (2/8) Garryaceae (1/18) Lennoaceae(3/8) (1 rarespecieson thedryCaribbean coastofnorthern Colombia) Theophrastaceae (5/110) (a fewspeciesofJacquinia and Clavija in SouthAmerica) Cactaceae(62/2.

LAURASIAN TAXA The phytogeographical ofthenewdevelopments significance inplatetectonics was first into focus brought byRavenandAxelrod thrusts (1974). ofthe whichtogether Orchidaceae. and Lennoaceae (also local in extreme northern ColomAlthough endemic.11 .000species of Neotropical epiphytes (Dodson. Maxillarinae. can be unambiguously referred either to theGondwanaland or to theLaurasian floras on accountoftheir distributional and thefossil general recpatterns ord. many including Malesherbiaceae.whichis one of the predominant taxa of canopytreesof Amazonianforests. 1980a). It is noteworthy thatmanyoftheendemic families listedin Table 1 are taxa ofdry.comm. Bromeliaceae Cactaceae dominate majorepiphyte families. subfamily Lecythidoideae (Lecythidaceae. 1979). bybothNorth ica. To thesemaybe added such specioseendemic groups as tribeBignonieae of Bignoniaceae (Gentry. Garryaceae. a listof29 Laurasianfamilies Similarly (or equivalents) in SouthAmerica whose majorarrival withbuilding probably coincided of the Isthmusof Panama in late Miocene to Pliocene was suggested. Koeberliniaceae (disjunct to Paraguay). Julianaceae. nine of them qualified as perhapsalreadyhaving had a priorpresencein SouthAmerica. manyof the Neo- families are ecologically tropical-restricted is one ofthe important. comprise nearly 5. becamegenerally platetectonics accepted. They listed51 basicallyGondwanaland plantfamilies (or equivalent units) that from SouthAmerica in themiddle probably to North America spread to late Cenozoic as the Isthmian barrier 54 families decreased.69 species each. and Loasaceae are predominantly meriaceae.and an additional (or whoseoriginal equivalents) presencein North Americawas probably olderbut whosemajorpresence there resulted from midto late-Cenozoic probably migrationfrom SouthAmerica. and amphicontinental KraCactaceae. or subtribes and Oncidinae Pleurothallidinae. Fouqueriaceae.pers. of areas. moreor less subtropical habitats. manydryregions. canopytreesof lowlandtropical forests. Martyniaceae.Familieslike and Vochysiaceaeare important Caryocaraceae. plants dry theseendemic taxamakean appreciable Although contribution to Neotropical speciesrichness.One ofthemajor oftheRaven-Axelrod was thefundamental synthesis difference between thefloras of the northern or Laurasiancontinents and thoseof thenow widelyseparated southern thatwere clustered continents together at the timeof origin of the Evenbefore angiosperms.whichis thepredominant groupof Neotropical lianas. iliesare morediversified ofthem. ecologically. thequestion ofwhy families evolved that intheNeotropics have moreprofusely speciated thantheir has notyetbeen Palaeotropical equivalents addressed. basically SouthAmerican fambia)-are dryarea specialists.566 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. Nolanaceae. ithadbeen realizedthatthe highaltitude SouthAmerican florawas largely derivedfrom north temperate sourcesand thatthe Panamanian isthmus had been a critical barrier to the southward of manynorthern migration taxa. One ofthecontributionsoftheRavenandAxelrod was pointing outtheremarkable synthesis degree to which many plant eventhoseshared and SouthAmerfamilies.). Unlike most of theirPalaeotropical counterparts. Prance & Mori. Humiriaceae. The onlyendemic tropical North American families-Crossosomataceae.

Although interchange it was less direct. themodern flora oftropical America with is ofremarkably bipolar ofitsplant families composition. perhaps such floristic in also occurred majorneotropical regions. myanalysis (Tables2-6) suggests adding together .389) (+4/361) (+3/120) (+ 1/6) (+4/515) (+16/1. Numbers indicate Neotropical genera in those genera(+ Neotropicalgeneraforwhichspecies estimates withknownspecies numbers/species are unavailable/total species in those genera). thePalaeotropics. having clearly that floristic elements one might theNeogenemixture oftwodifferent anticipate via the Central American isthmus could have had a majoreffect on increasing floristic almostdoubling theresultant flora ofeach ofthetwo diversity.594) (+ 15/1. Fromthisperspective.834) t = does not reach South America. * = listed as movingfromSouth America to NorthAmerica by Raven and Axelrod (1974). Laurasianfamilies as probableearlierarwere suggested otherpredominantly rivals in SouthAmerica.304) (+1/40) (+2/490) (+3/230) 416/5.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 567 TABLE 2. that ofLaurasian However.020) (+1/25) (+4/458) (+ 15/1.217) (+2/130) (+ 1/35) (+1/140) (+3/120) (+ 1/265) (+5/276) (+2/210) (+3/33) (+7/1. thegreat majority Laurasianor clearly Gondwanan affinities. tAceraceae Actinidiaceae *Aquifoliaceae Aristolochiaceae tBalsaminaceae *Basellaceae Bataceae Berberidaceae Betulaceae Boraginaceae (?) Buxaceae Callitrichaceae(?) Caprifoliaceae Caryophyllaceae *Celastraceae *Chloranthaceae Cistaceae Clethraceae Cornaceae Crassulaceae tCrossosomataceae Cruciferae Cyrillaceae Droseraceae Empetraceae Ephedraceae (?) Fagaceae tFouqueriaceae t(Garryaceae) Gentianaceae Geraniaceae tHamamelidaceae Hippocastanaceae Hydrophyllaceae tllliciaceae Juglandaceae 1/5 1/80 1/150 3/182 1/2 3/7 1/1 1/24 3/5 11/96 3/42 1/2 6/77 16/102 1/40 1/38 2/8 3/25 1/4 19/93 2/13 1/20 1/1 4/164 2/8 1/18 19/494 4/42 3/5 2/3 2/46 1/1 4/18 (Krameriaceae) Labiatae (?) tLennoaceae Liliaceae *Lythraceae (?) Magnoliaceae Myricaceae t(Nyssaceae) Oleaceae Orobanchaceae Papaveraceae tPinaceae Plantaginaceae tPlatanaceae Plumbaginaceae Polemoniaceae Primulaceae Pyrolaceae Rafflesiaceae Ranunculaceae *Rhamnaceae Rosaceae *Sabiaceae (?) Salicaceae Saxifragaceae Scrophulariaceae Staphyleaceae Styraceae Symplocaceae *Theaceae Theophrastaceae Typhaceae *Ulmaceae Umbelliferae Valerianaceae *Vitaceae Total 1/15 14/489 4/8 23/167 16/361 1/1 8/52 1/2 1/1 4/40 1/1 1/7 2/7 4/33 2/4 1/1 4/29 3/14 23/168 16/95 2/47 2/30 12/85 70/853 2/5 1/3 1/160 7/84 3/107 6/17 48/480 5/44 2/3 (+13/1.097) (+15/2.229 (+ 157/15. interrupted by largeexpansesof desertin anda persistent water barrier theSundaand Sahulshelves North between Africa in Australasia. As thusinterpreted. Laurasian elements of the Neotropical flora.279) (+5/730) (+4/185) (+1/30) (+ 1/130) (+2/130) (+ 1/10) (+2/110) (+ 1/200) (+3/505) (+ 1/450) (+14/1.

pers. 1982) has shown that a few South Americantaxa such as Dichapetalum.. Laetia.1973) (see Raven & Axelrod.).000m2sample in a South American Total: 106 species. 113 6 2 12 5 8 5 3 22 10 321 4 4 2 4 4 3 3 2 3 3 97 Dilleniaceae Euphorbiaceae Flacourtiaceae Hippocrateaceae Lauraceae Leguminosae Malpighiaceae Monimiaceae Moraceae Musaceae Myrtaceae Meispermaceae 2 7 Clsrca 2 1 1 1 5 3 2 2 1 8 1 7 1 3 1 Verbenaceae indet. and in prep.100 mm of rain 170 species over 2. Ind. would be expected (based on the regressionof species numbersversus precipitationcalculated fromthe 19 lower than the sites of Gentry. Gustavia. Ind. The Veracruz diversityis significantly expected inner tropicalvalue (R. sian?) genus. Fraxinus) Total 2 4 1 2 4 1 29 liana species. had alreadyreached Veracruz. Symphonia.5 cm diam. The Los Tuxtlas genus is Robinsonella. Cracraft. mammalianfauna (Patterson& Pascual. Veracruz (precipitation of 4. No. Graham (1976. Perozzi. Palmae Piperaceae Polygonaceae Rubiaceae Sapindaceae Sapotaceae Solanaceae Tiliaceae Urticaceaea Violaceae Total Laurasian Families Celastraceae Rhamniaceae S laceae Ulmaceae Total Unassigned Families Malvaceaeb No.000m2sample of lowland wet forestat Los Tuxtlas.568 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 69 TABLE 3. is especiallyprevalentin thetropicallowlands ofCentralAmerica. For a similar1. (cf. and the family may be fundamentally a b and Gondwanan elementshas not greatlyincreased overall Neotropical floristic diversity. 1981. 1966). Spp.). The northward Gondwanan taxa have so migrating overwhelmedthe corresponding southwardmigrating Laurasian taxa numerically that the latter's contributions to the total Neotropical florahave generallybeen This pattern relatively insignificant.whichmustonce have been populatedby a tropicalLaurasian floristic equivalentofthe endemicCentralAmericanherpetofauna (Savage. Summaryof transectdata forplants over 1" dbh in a 1.100 mm a year) (Gentry. 1982.g. an exclusively Central American (= tropical LauraLaurasian in origin. Gondwanan Families Acanthaceae Anacardiaceae Annonaceae Apocynaceae Araceae Araliaceae Bignoniaceae Bombacaceae Capparidaceae Caricaceae Compositae Convolvulaceae 1 1 2 3 3 1 4 1 1 1 2 1 4 2 2 5 7 4 1 2 5 5 3 2 9 1 2 2 3 1 10 2 8 9 12 7 3 24 7 3 1 No. including or southernCentral Americanforestwith4. 1972) or avifauna (e. Note the overwhelming preponderanceof Gondwana-derivedfamilies. Assignmentas "Gondwanan" tentative. Mexico by Miocene times. Spp. comm.). 1974: 625-626). prep. No.There are over 10 times as many Gondwanan-derivedas Laurasianderived Neotropical species. and Byttneria Yet the Paraje Solo palynoflorawas dominated by temperateNorth American .

189) 12/136 (+2/106) 44/801 8/125 17/142 (+ 1/30) 23/408 5/81 9/67 (+ 1/300) 13/87 52/1.Toledo (1982)has shownthatwithin Mexicotreespeciesrich- . Thus.Most of thisinvasion has been so recentthateven at the specific level therehas been little differentiation. individual tropical lowlandforests in northern Central Americamaybe less diversethantheirsouthern as suggested equivalents. Gentry.virtually all of the speciesthatreachnorthern Central America are infrom distinguishable SouthAmerican taxa (compare 1982cwith Gentry. Jarzen to emphasize moredirect prior.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 569 TABLE 4. between migration MexicoandtheGreater Antilles. As the Isthmian connection SouthAmerican closed.NumbersindicateNeotropicalgenera with known species numbers/neotropical species in those genera (+ Neotropical genera forwhich species estimatesare unavailable/total species in those genera). These taxa led Grahamand Engelhardtia. Anacardiaceae (?) Annonaceae Apocynaceae Bignoniaceae Bixaceae Bombacaceae Burseraceae (?) Caryocaraceae Chrysobalanaceae Cochlospermaceae Combretaceae Connaraceae Convolvulaceae (Dialypetalanthaceae) Dichapetalaceae Dilleniaceae (Duckeodendraceae) Ebenaceae Elaeocarpaceae Euphorbiaceae Flacourtiaceae Gnetaceae Hernandiaceae Hippocrateaceae Humiriaceae Icacinaceae 15/133 28/555 64/687 72/631 1/5 20/187 5/102 2/24 8/334 2/8 7/97 4/57 21/1. Amazonian-centerea Gondwanan families.of severalofthe although same north temperate generathattodayare disjunct in themidaltitude "bosque caducifolia" ofMexico:Liquidambar. by Sarukhan (1968).000 1/1 3/43 5/60 1/1 2/82 4/7 92/ general. Nyssa as wellas suchother Laurasiantaxa as groupslike tribes Tecomeae and Bignonieae of Bignoniaceae.there is a clearnorthward decreasein the number of species of manyNeotropical families (Gentry. 1969) was characterized by thepresence. Perhapsthisnorthward is-still migration taking place.980 (+48/8.607 28/267 1/6 3/22 12/114 8/46 13/56 (+2/257) (+3/250) (+2/125) (+2/120) (+ 1/100) (+2/125) (Lacistemaceae) Lauraceae Lecythidaceae Leguminosae Loganiaceae Malpighiaceae Meliaceae Menispermaceae Moraceae Myristicaceae Ochnaceae Olacaceae Palmae Polygalaceae Quiinaceae Rhizophoraceae Sapindaceae Sapotaceae Simaroubaceae Sterculiaceae (?) Tiliaceae Trigoniaceae Turneraceae (?) Violaceae Vochysiaceae Total 2/14 11/700 (+4/870) 11/275 216/2.866 (+88/12. Fagus. and Hauya. 1982a). At anyrate.110 (+3/42) 6/183 (+3/630) 4/53 5/24 27/438 (+5/490) 9/208 (+3/234) 11/106 14/293 (+2/360) 20/139 1/24 1/60 (+2/26) 11/98 (+2/650) 7/182 961/15. at reducedlevels. additional taxa moved north to completely dominate the Central American lowlands.It seems likelythat.Thereare onlyone speciesofTecomeaeand sevenofBignonieae in Guatemalathatare not also in Colombiaand Venezuela.904) elements. an Oligocenesite in PuertoRico (Graham Similarly & Jarzen. Nevertheless a number oftropical SouthAmerican taxa that must have arrived overwater werepresent. 1973).

69 Gondwanaland groups.545) Rubiaceae 1/2 Tovariaceae 2/92 Tropaeolaceae 7/88 (+6/653) Urticaceae ? 4/111 Zingiberaceae 1. dry 1978:75).266 Orchidaceae 1/8 Oxalidaceae (+2/870) 4/362 Passifloraceae 4/25 (+2/3.800) t = does not reach CentralAmerica.000m2sampleof richlowlandrainforest in a simiilar further south wouldbe expected (Table fewer vegetation speciesthan this is shown moist forest species:Mexican only bylowland 3).425/29.153 Melastomataceae 8/246 Monimiaceae 2/82 Musaceae 12/311 (+2/600) Myrsinaceae 27/277 (+3/160) Nyctaginaceae ? 306/8.254 1/11 1/18 14/275 1/37 5/8 3/92 1/1 7/43 66/1. America.Rzedowski.864 (+87/7.386 (3/356) Araliaceae 5/197 Balanophoraceae 7/15 Begoniaceae 1/600 Bromeliaceae 46/2.570 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL.218 (+10/1. Numbers indicate neotropicalgenera with TABLE 5. Sa.345 (+ 141/16.108 Brunelliaceae 1/51 (+7/712) Campanulaceae 9/568 Cannaceae 1/55 Caricaceae 3/29 Columelliaceae 1/4 Compositae 502/3.906 (+21/2. SouthernAndes/South Temperate 1/1 tAetoxicaceae 1/2 tAuraucariaceae Calyceraceae 4/46 Coriariaceae 1/1 (+ 1/170) Cunoniaceae 3/12 Cupressaceae 3/5 1/1 tEpacridaceae 1/1 tEucryphaceae 3/8 tFrankeniaceae 1/1 tGomortegaceae Hydnoraceae 1/6 (+ 1/100) Iridaceae 34/188 Juncaceae 6/49 2/3 tLardizabalaceae 16/592 (+ 1/15) Loranthaceae 10/270 (+ 1/11) Marantaceae 4/125 Marcgraviaceae 85/3. glaciation on thelowland of thetemlowering by the concomitant general mayhave been moreaffected inner taxa mayhave been eliminated or "contropical perature. Northern Andes Acanthaceae 61/1. Andean-centered species in those genera (+ Neotropical genera forwhich species knownspecies numbers/Neotropical estimatesare unavailable/total species in those genera). Whiledrought at themargin ofthetropics. condition American forests The relatively oflowland Central may depauperate and be due largely to Pleistocene climatic also have a muchmorerecentorigin ofPleistocene has been considered themajoreffect fluctuations.493 Araceae 38/1. ness of the lowlandtropical My in Veracruzalso show data from a 1. rainforest decreasesdramatically northward.861 1/1 (+2/1.000) Piperaceae 147/2. manysensitive .772) Total Loasaceae tMalesherbiaceae Myrtaceae tMyzodendraceae tNolanaceae *Onagraceae Podocarpaceae Portulacaceae Proteaceae tRestionaceae Santalaceae Solanaceae Winteraceae Total 12/266 1/27 24/1. pattern inbeing eveninultimately southern-derived areascontrast taxa.202) Cyclanthaceae 11/178 *Ericaceae 37/731 Gesneriaceae 40/917 (+3/590) Guttiferae 21/232 (+3/58) Haloragidaceae 1/1 Sb. very diverse. with manyendemic species(cf. * = listed as movingfromNorth America to South America by Raven and Axelrod (1974)..100) (+5/422) (+1/8) 199/4. However. Central tropics.

The most Laurasian clearly families to havenoteworthy ofspeciesinlowland complements SouthAmerican forests are Aristolochiaceae and Vitaceae.143 2/21 (+ 1/250) Xyridaceae Total 516/7. 1981).This mayhave been theultimate fateofthetropical North American flora thatis known to have inhabited even muchof the southern and central UnitedStatesduring theEocene and wouldbe consistent withsuchpatterns as themodemdiversity of Sabiaceae.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 571 TABLE 6.each witha single vinegenuswithnumerous speciesin SouthAmerica (Aristolochia. Threeof these-Buxaceae.Fourother tropical lowlandfamilies of probableLaurasianderivation are characterized by affinity fordryareas and a strong representation bothin theWestIndiesand in northern Central America. Boraginaceae.738) (+1/600) (+2/45) (+80/4. not botharealso wellrepresented surprisingly. intheWestIndies. comm. Miscellaneous taxa.799) fined southward" bytheslightly lowertemperatures during glacialadvances(Axelrod. and Rhamnaceaeare proportionately better in temperate represented North Americathanin the Neotropics. The northward movement of lowlandtropical Gondwanan elements has had no significant counterpart ofsouthward moving tropical Laurasian taxa. knownto have been widespread in North American Tertiary whichis greater in Panamaand Costa tropical floras.206 (+135/11.000 Capparidaceae 10/40 Erythroxylaceae 1/180 Koeberliniaceae 1/1 Martyniaceae 3/13 Velloziaceae 4/229 Zygophyllaceae 12/62 Total Unassigned Aizoaceae Alismataceae Amaranthaceae Amaryllidaceae Asclepiadaceae Butomaceae Canellaceae 91/2. Cissus).In their dryarea preference. these groupsare reminiscent of the . Numbers indicate neotropical with genera known speciesnumbers/Neotropical speciesin thosegenera (+ Neotropical genera forwhich speciesestimates are unavailable/total speciesin thosegenera). Rica thanit is in northern Central America (Gentry.525 2/4 2/61 7/188 26/799 46/932 2/7 3/11 Nymphaeaceae Lemnaceae Lentibulariaceae Linaceae ? Malvaceae Myoporaceae Najadaceae 4/12 3/116 3/20 50/860 1/1 1/8 5/27 2/5 (+1/230) (+4/416) (+4/90) (+7/535) (+3/210) (+3/280) Phytolaccaceae 13/78 (+2/38) Polygonaceae 10/203 (+5/724) Pontederiaceae 4/21 Potamogetonaceae 5/42 Rutaceae 36/233 (+4/313) Taccaceae (+ 1/30) Thymelaeaceae 7/71 (+ 1/10) Verbenaceae 40/1.). Guayana-centeredgroups Burmanniaceae Dipterocarpaceae Mayacaceae? Podostemataceae Rapateaceae? Saccifoliaceae Sarraceniaceae ? Tepuianthaceae Triuridaceae Total 13/51 1/1 1/9 19/151 15/79 1/1 1/6 1/1 4/12 56/311 (+4/416) Dry-area Gondwanan groups Chenopodiaceae 10/34 Commelinaceae 17/163 Cucurbitaceae 55/311 8/88 Cyperaceae Dioscoreaceae 1/15 Elatinaceae Eriocaulaceae 12/868 Goodeniaceae 1/1 Gramineae 127/838 Haemodoraceae 1/2 Hydrocharitaceae 8/20 Juncaginaceae Ceratophyllaceae 1/2 (2/14) (+ 18/3.692) Cactaceae 60/2.pers.

Even in SouthAmerica.and Ranunculaceae. Theaceae.Onlytwo genera(Cordia. intolowlandNeoancestry. theLaurasian-derived thereseemsto be a basic dichotomy between uplandand In Central floras. 69 Laurasianfamilies endemic tropical previously noted. and thusespecially well represented in thefossilrecord. One genus.Theophrastaceae has morerecorded speciesin Peruthanfarther north butis a morepredominant vegetational element and has in Central greater genericdiversity Americaand the WestIndies. Such knowledge of the of their recentarrival history of Andeanforests reliesalmosttotally on theworkof van derHammen invanderHammen.g. During thePleistocene thepalynoflora fluctuated . 1978) and restricted outcrops northern Venezuela(Styloceras goes southin theAndesbutmaynotbe closely related. perhaps southern). was uplifted thePliocene. thepenetration of thefamily is minimal and a few species of tropicalforests (monotypic Ampelozizyphus in drier itis better Gouania and Colubrina). forGondwanan-derived lowlandneotropical American upland withfamilies like Piests Laurasianelements clearlypredominate ecologically naceae. the principal By the beginning upheavalof the regionwas theearliest Pleistocene ofthis completed. Ulmus. which Theophrastaceae occursinlowland tropical butis poorly inAmazonia and mayhaveonlya single forests represented species reaching coastal Brazil. to Myrsinaceae. During glacialadvancethepalynoflora a primitive and depauperate such region suggested paramovegetation including as Aragoa (Scrophulariaceae).However. generaof other families mostJuglandaceae) reachonlyupland Panama. is minimal of Buxaceae. northern-derived elements ultimately Hypericum (Hypericaceae). Thefirst andhisassociates montane elements (summary 1974). (e.thepattern byRhamnus (Johnston & Johnston. SouthAmerican flora wouldbe almostimpercepWhilethelowland tropical ifall oftheseputatively Laurasian ofperhaps tibly changed tropical groups (a total SouthAmerica) wereeliminated. penetration of SouthAmerica to thedistribu(contrary in to a few limestone tion shownin Heywood.. representatives In thecase of all ofthesefamilies have penetrated intolowlandSouthAmerica. These ciallyimportant northern taxagradually decreasesouthward so that families likeHamamelidaceae and Pinaceae do notcrosstheRio San Juan lowlandsand are notpresent south of northern whileGarryaceae and manyimportant Nicaragua. Polylepis (Rosaceae.1981). Tournefortia) of thetwentythe lowland fourgeneraof Boraginaceaethatreach the Neotropics penetrate to any extent. Laurasianelements tendto prevailin montane forifnotalwaysin numbers ofspecies. Celastrus. intheCordillera toarrive atthePalynological sites Oriental attheSabanade Bogota as theCordillera wereHedyosmum andMyrica. Umbelliferae. in Neotropical Laurasiantaxa are muchmoreimportant montane floras.Conwind-pollinated thatthepalynological we maybe reasonably confident documentation sequently in SouthAmericais meaningful. Plantago. Gentry & Foster. a fewhundred speciesin all oflowland tropical In fact. Magnoliaceae. during of the Pleistocene. links Clavija.ecologically.572 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. and Ulmaceae espeas canopymembers of the temperate montane forests. Fagaceae. although represented partsofSouth America.Although tropical forests Rhamnaceaewere listedas basically Gondwanan shown byRavenandAxelrod (1974). Juglandaceae. 1978) seems typicalof the groupand pointsto a northern As in Boraginaceae. Valeriana.Manyofthesespeciesare ests.

000 appearedapproximately some southern Although in importance. Brazil/Peru westward. elements northern tocene. Juglandaceae.and (perhapssouthern) Styloproblematical taxa like Gunneraand phytogeographically withsouthern at the end of thelower arrived & Foster. Juglans. as Geranium. Note theconcentration Peruand adjacent Amazonian northern known in poorly collecting Additional frontier.Moraceaetribe distribution FIGURE1. families as Myricaceae. during and becamedominant Pleistocene Pleistocene Middle the of yearsago at theend 250.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 573 Fkxd NewIr opct ki b r r no 3 TROPICAL AMERICA Ldyer 1ft Md) of a taxonof canopytrees. enriched during weregradually floras forest Lysipomia. region diversity thehigh extend willprobably Colombia aboutby climatic brought ofthevegetation zonation altitudinal with thechanging periods. Gentiana.Alnusfirst ceras (see Gentry Quercusfirst the middlePleistocene. increased progressively and thereafter thePleistoOriental during in the Cordillera also arrived taxa like Weinmannia are forests Andean northern and the present elements prevailed cene.such additional along Urticaceaeappearin thepollenrecord.Tribaldistribution in thearea oftheColombia/ Amazonia western ofspeciesin (wetter) fig.1981). northern taxa. by Laurasian stilldominated Betulaceae. TypicalAmazon-centered thedata of Berg(1972: from plotted isohyets withspeciesdiversity Olmedieae.Boththeparamoand montane glacialand interglacial changesbetween In thelowerPleisthePleistocene. Even todaysuchnorthern .1).

to be Predominantly herbaceous Laurasianfamilies have a greater tendency . Magnoliaceae. and Caprifoliaceac are present in tropical South Americaalmostentirely in the uplandAndes. Cornaceae. theabsolutedominant ofmostColombian lowermontane forests. white sand substrates (manyTernstroemiaand Ophiocaryon). or secondgrowth (Trema. and Staphyleaceae.Theaceae. Celtis). theonlyAmazonian speciesof southtemperate Podocarpus is restricted to whitesand (Gentry et al.Oleaceae. Interestingly. apparently an undescribed species). Cyrillaceac. hlexinundata. Clethraceae. Ophiocaryon duckei. Other woodyfamilies like Salicaceae.thereis a decrease in representation of thesefamilies farther south. Ophiocaryon. hlex. Ampelozizyphus(Rhamnaceae)). 2887] (MO) fromnear Iquitos.Berberidaceae.Turpiniaoccidentalis).Similarly. Ulmaceae. Sabiaceae. thelowlandrepresentatives of suchtaxa are often restricted in Amazonia to ecologically impoverished extreme sitessuchas seasonally inundated streamsides (Salix.Aquifoliaceae. Hippocastanaceac.For example. Within theAndes.Quercus.69 O~~A>D _ _ l _ 3C_ _ 0-Lt _I_ A 97 .574 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL.Trema mnicrantha and Celtis iguanea. Ternstroemia. Celastraceae. have one or two widespreadspecies(or genera) thathavebecomewidespread in thetropical lowlands (respectively: Salix humboldtiana. Gouepia. nb f i r i ) Fagaceae. does notoccur in Ecuador.

Extra-Amazonian Smith 213). la Vriesea - - ~ - h Tillondsia > _ _ Vriesea ~~~~~~~~ ~~Catopsis ~ ~ ~ ~ rise t ~~~~~~~~~~~Vriesea \ vet~~~~~~~~~~~~~~ Tilan'i I~~~~~~~~~~~ ~~\s a f t ~~~~~~Tillandsia taxepiphytic ofa predominantly distribution orAndean-centered FIGURE 3. subfamily .i \ ~~~~Guzmania .\iltondito .1977:fig.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 575 SUEZ C CCOtXCT-00 S. & Downs. Tillandsioideae. on (Bromeliaceae.

Guayana andassociated as part ofAmazonia except forspeciesnotfound elsewhere inAmazonia). 9 forcomplete FIGURE 4. Southern Andes. Ranunculaceae. Northern Andes. Northern Venezuela-Colombia. C = Canopy treesand lianas:percent cutrngin that ofregion's speciesofthat habit group which are endemic to habit occurring in each region (percent andpalmettos: percent ofthespeciesoftaxacharacterized theregion inparentheses). 3. to theregion in parentheses). Nevertheless. weeds. Orobanchaceae. 7.6. (western dies. regions.117recently monographed speciesocor endemic to each region.s. HyBalsaminaceae. Guianasubregion (included border defined by500mcontour). Rosaceae (s. Saxifragaceae. 9. better represented in themontane Neotropics thanin thelowlands. and Plantaginaceae are much drophyllaceae. E = Epiphytes which are endemic by habit occurring in eachregion (percent ofregion's speciesofthat habit group data. Gentianaceae. 8. Cerrado speciesoccurring dryareas. 2.576 GARDEN ANNALS OF THE MISSOURI BOTANICAL 69 [VOL. Mexico andCentral America. Amazonia Highlands (over500in). T = Percent ofthespeciesoftaxacharacterized by region.Umbelliferae. Polemoniaceae.5. Scrophulariaceae. See Tables7. Cruciferae. 8.Callitrichaceae. 1. is a strong dichotomy between thenoticeable For bothtreesand herbsthere absence in the presenceof Laurasiantaxa in montane forests and their virtual lowlands.families likeCrassulaceae. 4.Smallnumbers indicate percent of monographed in oftotalsampleof8. Caryophyllaceae.). West InNeotropical phytogeographic exceptfora few weedyand their patterns are notso wellmarked. Primulaceae. 10. CoastalBrazil. Plumbaginaceae. Geraniaceae. .

especially MostoftheLaurasian inpart their recent this reflects 1982a).Theyare well Andean and only5% intheSouthern region in thecoastalBrazilregion constitutes species). families SouthAmerican within fortheseautochthonous are centered groups. graphed Neotropical reother FloraNeotropica-caliber butalso including as many pica monographs. and virtually noneofthatofthelowland tropical flora GONDWANAN TAXA floraare Gondwanaland-derived of the Neotropical The majorcomponents difference phytogeographic there is a fundamental groups (Tables4-6). 2.Manyof thesefamilies tently intotwomajordistributional to as Amazonian-centered. AMAZONIAN-CENTERED TAXA of the totaldata set of monotaxa include38%o The Amazonian-centered have an averageof almosthalf(44%) of their graphed species.on theaverageonly12%oftheir speciesoccur under-represented inthenorthern Andes.especially maindiversity taxa although their be termed Andean-centered mayconveniently elevation inthelowlands andinmiddle nearthebase ofthemountains is reached rather thanin thehigh mountains themselves. Thesetaxa thenorthern tersintheAndes. These families all ofthesetaxa are predominantly canopytrees speciesin Amazonia.715spp.) or clearlynorthern-Andean-centered clearlyAmazonian-centered families of each of thesemajorphytogeographical (2. however.moreover. almostall canopytreesand lianasoftheNeotropical lowThesetaxaare conspicuously landsbelongto theseAmazonian-centered groups. journalsforthe botany in majorsystematic visionsof majortaxa as I couldfind a data set of 8.117monographed species.This yielded shownin FigtheNeotropics intotheninephytogeographic regions Subdividing in each region speciesoccurring ure4.Virtually and lianas. treesandlianas. secondary monographed majortaxa has as manyspeciesin coastalBrazilas in Amazonia.). .Presumably little intheAndes(see Gentry. last tento twenty years. beginning with theFloraNeotrofamilies and largegenera. 3). ilies in Table 2 accounts for a very small percentage (<10%o) of the total Neo- tropics.In orderto becameaware of thesetwo striking I extracted dataforrecently monoand compiled distributional document them. Figs. The greatmajority (71% or provided the data summarized thatare either speciesbelongto families almost5. Fig. 1) and maybe referred andcontrastingly with extra-Amazonian arefundamentally Mostoftheremainder cendistributional in Amazoniaand usuallyhave their verypoorrepresentation Andes(Table 5. and counting thenumber ofmonographed in Tables 7 and 8. fallconsisthisgroup. epiphytes.which (16% oftheir represented one ofthese nota single a distinct center formostofthem. However. in theAndes. canopy and palmettos. theimpressive arrival. in Amazonia(Table 4.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 577 rather thewoodyones.052 spp. cloudforests I first patterns in thefield.theextra-Amazonian ones are chiefly shrubs. The component butalso amazdistributional patterns notonlyshowstrikingly concordant groups taxa are overwhelmingly The Amazonian-centered ingconsistency ecologically.have speciated taxa. famlistof72 Laurasian-derived Neotropical As a result.800species)of monographed (3.

829 (1. Andes 723(481) 128(51) 126(73) 71 (47) 373 (154) South.390) 1.567/8. 4 Betulaceae.288 (940) 932 (676) 1. Proteaceae.334(1. aquatics.263 (683) 1.934(1. Habit' Epiphytes andpalmettos Herbsand shrubs Vines3 Canopy trees and lianas Amazonia 1. Polylepis. 5 Parasites.117 in region occurring 17% 12% 8% 8% 14% 'Predominant habit ofmonographed taxon. Juglandaceae.072) Coastal Brazil Cerrado & Caatinga North. Brunelliaceae. Clethraceae. . 2 Figures forGuianassubregion include onlythosespeciesthat occurin thelowland butnot Guianas Distribution primarily reflects pattern ofsingle largefamily-Passifloraceae (363spp. Colu Rhamnus.).TABLE 7. many Mexican desert shrubs. Number ofspeciesandendemism inmonographed taxabypredominant habit group andg speciesforeachregion inparentheses.165) 23% 76% 16% 73% 11% 73% 18% 56% 10%70 16% 54% 24% 60%o Percent ofspecies % Endemism6 (total= Totalendemism7 = 81%) 6. 6 Endemic speciesin region as percent oftotalmonographed speciesin that region. Andes 469(231) 437(231) 78 (40) 47 (37) 31(20) 57 (39) 144 (85) Central America & Mexico 453 (189) Aridarea trees Miscellaneous5 Total (8.117) Montane trees4 292(205) 60 (16) 75 (40) 7 (5) 498 (426) 184(140) 140(76) 167(103) 74 (40) 45 (18) 2 (1) 80 (67) 12 (8) 482 (322) 373 (272) 559 (319) 607(443) 132(65) 98 (84) 61(52) *0 17 (12) 143(130) 3 (1) 4 (1) 29 (12) 37 (30) 48 (35) 1. Podocarpaceae. 7 Endemic speciesin region as percent oftotalspeciesin all monographed taxa.454 (819) 1. Velloziaceae.

052 2. Podocarpaceae.. Brunelliaceae. largefamily-Passifloraceae ofsingle patterns reflects primarily 4Distribution Colubrina Rhamnus. N. 3.) 256 105 3. Polylepis. Clethraceae. V zue & Mexico Colo 44 11 6 16 16 18 14 16 trees5 Montane Others (parasities. & Caatinga Andes Amaz. Braz. shrubs. aquatics. Percent Cerrado North. Habit' treesandlianas Canopy andpalmettos Epiphytes Herbsand shrubs Vines4 Arid area trees Coast.). Proteac.117 Total I Predominant taxon. Total Spp. ofmonographed habit butnotalso i Guianas occurin thelowland include Guianassubregion onlythosespeciesthat 2Figures for occu in which in eachregion they sincewidespread to morethan100%o 3Percents total speciesare counted (363spp.034 459 South. desert Mexican many . Juglandaceae. Andes Percentof Species3 5 17 42 45 Central Amer. 5 2 10 12 7 16 16 12 27 12 27 4 1 28 27 17 15 22 54 29 1 38 35 496 12 16 29 6 6 10 8. 5Betulaceae.715 1. Velloz.TABLE distrib andgeographical group habit taxabypredominant ofmonographed distribution 8.

Yet mostother Central American Bignoniaceae areundifferentiated from their SouthAmerican progenitors evenat thespecieslevel. in almosteverycase the cerradospecies are shrubsor subshrubs in taxa thatare otherwise treesor lianas. therelatively fewAmazonian-centered Nevertheless.forspecific examplesof these patterns). An interesting subsidiary pattern is shownin Central America by severalof thesetaxa.andnonecrossing theHoldridge system tropical/subtropical delimitation at 12'N latitude in Nicaragua(exactlythe same latitude as the diversity-reducing Isthmus of Kra in Malaysia!). Manyof theAmazonian-centered families have derivative speciesin thecerrado and associateddryareas oftheBrazilian shield (12% of the species). may be referred to conveniently as Andean-centeredand is almost .Most of thespeciesof thesetaxa thatdo reachCentral America are notendemic. canopy(Table 3 and Gentry. Severalofthegroups have a distinct secondary radiation in northern A good exampleis provided Central America.000monographed Thus.whose strong representation in the coastal regionwas emphasized by Kubitzki (1975). 1982a. taxa thatreach northern Central America continue to constitute virtually all ofthelowland forest in prep.Crescentieae thatgroupin such important features as indehiscent fruits andbat-pollinated flowers that have sometimes they beentreated as a distinct family (see Gentry. The Amazonian-centered taxaarepoorly represented inCentral America (only 15% oftheir species). generic. Thiscontrasts with in Amazonia. theyare mostly thosefewAmazonian speciesthathappen to have unusually wideranges. Most of thesefamilies have severalspeciesreaching eastern fewer Panama. Within Central America there is a marked decreaseintherepresentation of Amazonian-centered taxa from southto north.580 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL.where35% of all the species of these taxa are 80%o endemism endemic.).These groups are markedly poor in the WestIndies(10%o of the species on the average. ANDEAN-CENTERED TAXA The second major Neotropical phytogeographic pattern. differentiation.only6%o species ofAmazonian-centered taxaare endemic to Central America. just as in Amazonia.which is rather sincethey surprising makeup nearly all of the moistand wet forest canopyof the Central American lowlands. ofthe3.Those Amazonian-centered familiesthatdo extend farther intoCentral America typically have onlyone or two species north of Nicaragua(see Gentry.Clearly reflect a longhistory of differentiation in Central Americasubsequent to an initial colonization by SouthAmerican Tecomeaestock. by Bignoniaceae with tribe Crescentieae three and35 speciesalmost having genera inCentral America exclusively derived (Gentry.1980a). It is tempting to think ofsuchpatterns as reflecting a twopulsemigration: (1) earlycolonization across theproto-Antilles by islandhopping at theend of the Cretaceous with subsequent majordifferentiation and (2) a majormigration subto closing-of sequent theIsthmus ofPanamathatwas too recent to permit much or even specific. rather.contrastingly extraAmazonian. 1979.witha disproportionate partofthattotaldue to theevolutionary explosion of a singleotherwise smallsectionof Tabebuia). 69 not even Dilleniaceae.Although from thefundamentally SouthAmerican tribe are so distinct from Tecomeae.noticeably western Panreaching ama and CostaRica. Crescentieae 1974a).

are also better in drythanin wetareas. especiallyCosta Rica and Panama.and are also wellrepresented in thesouthern Andes(17% oftheir species) (Table 8). theyare well represented in thecoastalBrazilregion (18% oftheir species)and poorly representedin northern Venezuelaand theWestIndies. account forthegreat majority (71% ofmysample)ofNeotropical plantspecies.theyare also poorlyrepresented in the drycerrado-caatinga region (7% of their species). The Andean-centered taxashowvery pronounced inCentral endemism with America. where22% of their speciesoccur.Gesneriaceae. DRY-AREA-CENTERED TAXA A fewsubsidiary distributional need to be mentioned.Together thesefamilies. Ericaceae. Caricaceae.Cyclanthaceae. Dry-area-centered represented taxa tendto be largely shrubs and herbsalthough somewellknown treegenera likeProsopis . understory shrubs (Acanthaceae. thathighspeciesdiversities do not occurat high altitudes butrather inthewetlowland andpremontane cloudforests alongthebase and lowerslopesofthemountains. Thesetwodominant phytogeographic patterns-Amazonian-centered trees and lianasand Andean-centered and epiphytes-together palmettos. latter often specialized forthehighly alkalineconditions typical of deserts. Monimiaceae.Solanaceae).Orchidaceae.Melastomataceae.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 581 themirror imageofthat shown bytheAmazonian-centered taxa. Marantaceae.). Threegood examplesare Capparidaceae. bothCentral Clearly America and western SouthAmerica havebeen majorevoforthesegroups. Piperaceae.Araliaceae. and coarse palmetto-type monocots(Musaceae. These groupsare notonlyconspicuously under-represented in Amazonia(11% of theirspecies). lutionary centers of thesetaxa is highest Although representation in mountainous phytogeographic it shouldbe re-emphasized regions. absolutely dominate thelowlandneotropical bothin Central flora. Piperaceae/Peperomia. Familieswiththispattern have their distributional centers inthenorthern Andes.In thoseregions where Amazonian-centered taxa are wellrepresented.where overa fourth (27%) oftheir species occur.thismaybe mostly ofthemuchpoorer floristic data base from northwest SouthAmerica. Southern Central America is clearly a major ofspeciation secondary center formostofthesegroups.Thus any explanation of the ofevolutionary patterns diversification in thesetaxa willlargely therichexplain ness oftheNeotropical flora. verywell represented in Central America. thesegroups account for most of the incredible floristic of the Choco region(Gentry.Bromeliaceae. contrast to the low (42%) Central American endemism of Amazonian-centered taxa (Table 9). and SouthAmerica. diversity 1982a). all basicallyGondwanan. someofthese Although groups actually appearto have morespeciesin Costa Rica or Panamathaninthe northern an artifact Andes. etc. Zingiberaceae). Myrsinaceae. Rubiaceae. Like theAmazonian-centered group. In anyevent. shrubs. Amaranthaceae and possibly the Chenopodiaceae. Unliketheir Amazonian-centered theAndean-centered taxaare counterparts. Guttiferae. These groups are predominantly epiphytic (Araceae. and Zygophyllaceae. 73% of the Central American This is in strong speciesendemic. One is that of patterns taxa withdistributional centersin dryareas and poor representation both in Amazoniaand themoistAndes. Cactaceae. Andean-centered taxa are poorlyrepresented and vice versa.

Andes 41 67 40 58 66 25 South.TABLE in eac ofthosespeciesoccurring Percent regions. ofphytogeographic endemism 9. Relative Total% Endemic to I Reg. ofmonographed habit 'Predominant (363 spp. largefamily-Passifloraceae ofsingle patterns reflects primarily Distribution Rhamnus. Clethraceae.). Colu 3 Betulaceae. Juglandaceae. Brazil 67 86 54 54 67 50 Cerrado & Caatinga 73 76 62 40 33 71 Ce North. 81 81 82 71 85 84 Habit' andlianas trees Canopy andpalmettos Epiphytes Herbsand shrubs Vines2 Montane trees3 Arid area trees 2 Amaz. Polylepis. Brunelliaceae. desert Mexican many . shrubs. 80 70 27 53 71 - Coast. Podocarpaceae. Andes 59 49 53 51 68 79 A M taxon.

1978)has pointed region in thecerrado area taxa even though species of in is most among dry striking mism Mexico numin absolute of terms flora country's the dominate forests lowlandtropical from the south strongly derived taxa ultimately the endemism. the region's America (57% 53% Central out thatende(62%). predominate 1973).rather Mexican seemto be autochthonously ofthesedryarea taxamight many although of species there. just as Rzedowski speciesrichness. species). 1963).hereincluding (42% oftheir Andeanregion southern Mexico and (54% of their species). ilieslikeFouqueriaceae. Even some amphitropical longdisrecent from relatively believedto result likeLarrea are now generally 1976). than distributions rather ancient tancedispersal basis. Rzedowski(1962. of the Brazilian region drycerrado-chaco-caatinga in theSouthern Anof thesetaxa is aboutas strong representation Although in is greater endemism slightly American region. .Not surprisingly. restricted edaphically originally times. and seasonally outcrops ofdryarea disjunctions range although amphitropical It should be notedthat.Crossosomataceae..Such pat(Rzedowski. recentlong distancedispersal manyof these surelyreflect taxa are frequent.are all primecandidates betweenChile and California. bers. Rzedowski. Cactaceaeindry theuninterrupted ofdrytaxa. due to ancient thannecessarily forlong forexample.1978)as evidence areas.implying persistence history ofa longevolutionary that (1979)suggests mostoftheCenozoic. Cretaceous as limestone forsuchedaphically are specialized drysubstrates whosemembers inundated savannahs).g.especially American region and the Central taxa adaptedto dryareas are also America. and Malesherbiaceae. oftheUnitedStatesdeserts dryarea flora derived famendemic ofwell-marked ofa preponderance theprevalence especially terns.Whether and Quaternary Tertiary in Mexicoandnorthern Central ofmany ofthesegroups differentiation thestrong via late Americaimpliesthatsome of theirancestorsmay well have arrived tribe Crescentieae above. Bignoniaceae.havebeencited(e. shrubs dry-area distancedispersal(Carlquist.Despite high in contrast to thenorth temperatein theMexicandryarea flora. 1962.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 583 in the These groupsare best represented and Bulnesia also show thispattern. regimes dryclimatic to theincreasingly Generadisjunct or autochthonous arrival origin. partofthemonte. based on their American preponderance Central and northern so by affinities or presumably clearlyof Gondwanan mostof themare either of dryarea adapted analogy. above. Moreover. on an individual be interpreted ofdryarea plantsmust rangedisjunctions of the and interesting component Whilethesedryarea taxa are a significant in terms ofoverall Neotropical are relatively unimportant flora. Central northern andthe region in thenorthern Venezuelan-Colombian wellrepresented relatively shield. emphasized thanthe ancientpatterns (Raven. mostof islandhopping (cf.Clearly (Wells& Hunziker. Axelrod ofdryareas at leastthrough of dryarea taxa mayhave been and differentiation muchof theearlyevolution in thelate expanded as dryclimates taxa spreading dryareaswith in edaphically or not. Lennoaceae.Thus the highspecies numbers phytogeographic a secmostly are probably shrub and herbtaxa in Mexico and adjacentregions inresponse diversification from activeevolutionary resulting ondary phenomenon rather zof thePlioceneand Pleistocene. as in theCentral dean region more and even pronounced of species) vs.1982). they Neotropical (1962)notedforMexico.

versification.. opposing araimeais closertoTiliaceae(Kostermans. Finallythe GuianaRegion. andDipterocarpaceae & Ashton. 1970).1974).comm.boththerecently Andesand mostofAmazonia. Bambuseae (Soderstrom & Calderon.584 ANNALS OF THE MISSOURI BOTANICAL GARDEN MISCELLANEOUS SUBSIDIARY [VOL.1974)and some of thearchaictaxa of coastalBrazil-e.and especially the GuayanaHighlands. a Cecropiawiththesimply spicatefemale inflorescence ofAfrican Musanga (Berg. whichare in turn no morethana northern phytosubsetof Amazonia.Moreintriguing are several non-endemic that arefound These includeSarraceniaceae Neotropics onlyin theGuayanaregion. are well known as areas ofhighendemism and muchphytogeographic interest (Maguire.)-maydatefrom theCretaceousseparation of SouthAmerica and Africa. Nevertheless. region The floristic termed matter are drawn). ofthespeciesofthesummits Steyermark.pers.Even many are sharedwiththelowlands.Coastal Brazilis noteworthy fortheconcentration of often primitive speciesin a restricted area (e.g.Geologically thisarea is veryold. uplifted which was underwaterintothe Pleistocene. Tetrameristaceae theother (twomonotypic genera. 69 PATTERNS A fewother Neotropical phytogeographical patterns merit specialnote.. view that the Pak(see Maguire 1978. in Chocoother than as partoftheNorthern ofdistribution itschief center Andean region. Kubitzki. Howbetter represented either ever. Nevertheless no family has its distributional center incoastalBrazil.To be sure.Perianthomega (intermediate betweenthe two maintribesof Bignoniaceae and perhapsclose to the ancestral stockof theneotropical Bignonieae).andtheplants ofthetepuisummits have had the potential forverylongperiodsof evolution in isolation. between summit flora exchange andthelowland forest flora that ascendsthetepui slopes has apparently been muchmoreextensive thanonce thought (compare and Maguire. A veryfewsmallfamilies do have their ofNeotropical centers diversification in theGuayanaarea. and theapparent ofunspecialized taxa in CoastalBrazilmaysuggest prevalence theimportance ofthis as a sourceareaforother region phytogeographic regions.Thisrich. subsetof whatis here young. 1979. (disjunct from North inAsia). theNorthern significance oftheChoco area is almost entirely at thespecieslevelalthough itdoes have a fewendemic genera likeTrianaeopiper andCremosperma (Piperaceae) No family has (Gesneriaceae). and 1978)is based on weakevidence is phytogeographically irrelevant sincetheSouthAmerican taxonclearly belongs to theDipterocarpaceae ancestral wherethetaxonomic limits plexus.Such patterns ancient notactiveevolutionary disuggest survivals. 1975). primitive Dilleniaceae(Kubitzki. 1970).thereare a few strikingly geographic distinct be recognized endemicspecies and generain the regionthatmight as distinct families-Saccifoliaceae and Tepuianthaceae (close to Gentianaceae). 1975. are relatively recententities geologically speaking.The samefamilies that arewellrepresented thereare invariably in Amazoniaor theAndes. America). 1982a) is theChocoregion ofPacific coastalColombia and adjacentEcuador. butgeologically is an important perhumid. Another rather isolated lowlandarea notedforitsendemism (Gentry.g. The onlygenerally accepted families that seemto showthis .Soderstrom & Calderon. (close to families inthe Rutaceae).

FLORISTIC SUMMARY To summarize Neotropical floristic patterns.The second. we havetwomajordichotomies. secondimportant in theAndes.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 585 pattern are Burmanniaceae. andpalmettos mostly to Gondwanan with belong northern groups Andeandistributional centers.It is now Andes.especially centers groupthathas distributional thenorthern in Amazonia. even a fewendemic speciesbecomenoteworthy. Mayacaceae. withrespectively 9% and 8% of thetotalof monographed species.somecenters ofendemism based on as fewas two species. ofclearly Laurasianfamilies to extensively invadethelowlandNeotropics are vines(Arisin forest tolochia.Gouania) mainly occurring edgesituations .Together theyaccountfora totalof not morethan some 300 species.has notpreviously beengenerally saprophytes. Thurniaceae. rather than Cissus. Two other regions that inplantspeciesas judged are surprisingly depauperate from this data set are the West Indies and the northern Venezuela/Colombia region. muchless thanthe 90Wo although suggested by Maguire(1970). thetotalflora ofthatregion is minusculeindeed.Epiphytes. all lianas and canopytreesof the lowlandNeotropics Virtually belongto Amazonian-centered taxa. Podostemaceae. theoverall floristic significance oftheGuaand yana area. likeGondwana-derived herbtaxa they have extra-Amazonian distributional patterns. In general aquatics. evenwhenallowance is madefor therelatively small area ofupland Guayana. Venezuela/ forany of thesephytogeographical Colombiais by farthe lowest such figure In thislight it is clearhow Steyermark regions.and tendsto be poorly represented clear thatthe Laurasian/Gondwanan results from of dichotomy the separation North and SouthAmerica mostof Cenozoic time. understory shrubs. and betweena largegroupof families withAmazoniandistributional and a centers. Many herbsare from Neotropical widespread predominantly north temperate groups. The 77% endemic speciesof theGuayanaHighlands is slightly higher thanthe similar figure forany otherphytogeographic region (Table 7).On theother endemism oftheGuayana hand. speciesthat occur in theGuayanaHighlands is anyindication. Vines (as opposedto lianas)are represented in bothLaurasianand Gondwanangroups. Canopy trees of montane forests come fromboth Laurasianand Gondwanan with a gradation in therelative groups of importance thetwofrom north to south. it is noteworthy that theonlyrepresentatives however. All are specializedforunusuallife styles. although equallyclearcut. is somewhat supported by mydata set. Withsuch low totalendemism.or semiaquatics. (1979)was able to achievesuch a finescale in delimiting centers in Venezuela. Triuridaceae.within thepredominant latter group.thefamed high highlands.The fundamental dithrough between and Northern chotomy AmazonianAndean-centered families. now seems verymuchless thanearlier Ifthe3% ofmydatasetofmonographed believed. The first is betweenthe basicallyLaurasianmontane floraand basicallyGondwananlowlandflora. The value fortheWestIndiesis almostidentical 59% endemism to the60%o overall ofCentral butthe24% figure for endemism northern specific America. especially the GuayanaHighlands.

1982)). 1978).Frankie (Frankie Leguminosae et al. Acanthaceae.Scrophulariaceae Labiatae. logically forexin some groups.oftenlarge.. even in the Andes wheretheyare ecospeciatedverylittle Americatheseplantshave producedrelaEven in Central dominant.g. tubularflowers bees mostly belongto Amaby specializedlargeand medium-sized pollinated 1974a.many PassiOrchidaceae..Almost differential One striking none of the whereasvirtually trees are wind-pollinated of Laurasian-derived groups Andean-centered is. regions and amongthedifferent in different patterns speciation Laurasian. hummingbird-pollinated or evenpredominantly. groups is rather uncommon system ceae) whereasthispollination Polemoniaceae. groups in all phytogeographical wellrepresented probliesintheir groups phytogeographical/habit A related between difference have wind-pollinated.). Manyofthenorthern families angiosperm Gondwanan (e.Campanulaceae-Lobelioideae.Lecythidaceae (Frankie Cochlospermaceae et al. 1983. (e. Lythraceae. zonian-centered (Prance.and LeguminosaeamongAmazonianaceae.g. Ecotypicdifferentiation tively sets of related speciestendto ampleamongoaks in Costa Rica wherediff&rent 1977. al. systems otherpollination Taxa having mostly specieswithsmall flowers). Tropaeolaceae. Musaceae. & Baker.1976. Marantaceae. largely upland.The lifezones (Burger. 1974. families (Apocynaceae. thaceae-Loranthoideae. flowers (sensu generalist-pollinated and showno obvioustrends. pollinator species. inconspicuous generahave tiny are et 1982b) Frankie Gentry.Bignoniaceae(Gentry.Mori & Baker.. Gesneriaceae. in SouthAmerica.ZingiberaMarcgraviaceae. (1) between are thedifferences noteworthy Especially thelatter groupsand (2) within Gondwanan. in other (e. WoodyLaurasiantaxa. amongLaurasiantaxa. Plants withratherconspicuous. Ericaceae. with representatives in thetemperate zone tendto have Neotropical pollinated mostof whose Neotropical (e.g. groups amongother butare also found large-bee that mostly are Laurasian groups Even some Zingiberaceae). mature AmerCentral bothin northern distributional centers with pattern amphitropical Andeanregion. based on thelonggeneration Central of mostof the taxa in SouthAmericaand even southern arrival recent ofplanttheimportance that with models emphasize It is also consistent America.. some miscellaneous Convolvulaceae.g. taxa and lowland. Combretaceae. ica and thesouthern EVOLUTIONARY IMPLICATIONS different radically suggest patterns phytogeographic These verydistinctive adaptivetypes. floraceae.. is frequent few species.predominantly groups.. Holdridge to different be restricted withexpectations in thesetaxa is consistent of little speciation pattern general coupledwiththerelatively timesofwoodyplants. a few Gentianaceae. unavailable in promoting a potential patently interactions speciation.586 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. Bignonispecies of Apocynaceae.pers. 69 show an in dryareas and typically Shrubsare best represented forest.LoranBromeliaceae. are largely.1974). andAndean-centered Amazonian-centered thepredominant between group halfof thefamilies is in pollination ecology.. 1974. to wind-pollinated that canopytreesand lianasare thegroups The woodyAmazonian-centered withclassicalzoologicalmodelsof alloconsistent patterns showbiogeographic . centeredtaxa). 1974b). largely able modesof speciation. Scrophulariaceae.comm.

In general. zones. and palmettos understory The epiphytes./genus a regudiversity perhapsapproaching over 100species)and community having 1982b).hybridization (e.notedsimilar high("palaeopolyare frequently numbers therule(Bawa.g.. sia. by whatappearsto have been taxa are characterized Andean-centered northern muchofit sympatalmostcertainly and adaptive radiation. (Wright.g.Berry Luteyn. Psychotria.g. and Dichapetalaceae) Chrysobalanaceae. distributions-alloof restricted patterns thekindsof correlated to demonstrate fit groups-that in unrelated patric amongcloselyrelatedspeciesbutreplicated It is forest speciation. pers.Templeton. theme (Gentry. very many local endemics equivocalwith model weredistinctly refuge Pleistocene veritable"species the regionand constituting scattered throughout occurring Piper. Marcgraviaceae. pollination specific conditions. well as their timesof theseherbaceous forrapidevoshouldprovideideal conditions relationships. refugia. 1974.). phenomena bothby the smalllocalized or genetic transilience 1980)optimized (Templeton.19821 GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 587 that have ofdistributions havethekinds Thesearethetaxathat patric speciation. Microgeographic.1979. (Gentry. 1970).g. 1982d. taxa (Caryocaraceae. lutionary as suchphenomena in thesegroups involves frequently evidencethatspeciation (e. refuge modeloftropical thepredictions ofthePleistocene withtaxa of not surprising thatForero(1976) and Lleras (1978). latedecologicalequilibrium thatmakeup mostof the shrubs.and Cavswarms"in largeevolutionarily plasticgeneralikeAnthurium.chromosome 1980. ofthehigh oftypical much diversity ciation couldexplain to fitthe Choco florato the expectations of the 1982a) attempt my (Gentry. Berry.). Major shifts sand "campinarana" habitats as white in suchmarginal to specialization leading is a majorevoforests or inundated "varzea" or "tahuampa" forests seasonally species closely related 1982e). polyploidy Bedell. 1974). taxa(e. 1980b.)suggested 1983).Berry.Ehren& Gentry. which andseparating landslides.1977. Thereis some even understableclimatic differentiation.In thesetaxa outcrossing patterns woodylianas.and hybridization dorfer.Smith centered hisanalysis ofspeciation from (1980andinprep. (Goldblatt stablein a genusorfamily ploidy")and often 1970. lutionary withrelaseems somehoworderly distributions and speciation show allopatric on the averagewithonly19 genera tively fewspeciespergenus(14 spp.The samepatterns & Downs.. 1969).andfrequent vegetational callycoalescing The relatively shortgeneration provideopen areas forcolonization. 1958. regularly rather or shrubby typically groups. (e.. balance" in thisgroup reflect "shifting might in Fuchsia thatspeciation patterns reorganizations 1980)withmajorgenetic 1977. Harling. pers. of a habitat by partitioned and theneed forconstant recolonization populations climatic shifting cyclilocal rainshadows and other effects.1980)or cleistogamy is probably speciation perhapseven moreor less sympatric. explosivespeciation large(20 speciespergenuson theaverage. 1980. in mode of pollination may be rare while speciation least 120 ric. Lecythidaceae.. .Ashis rareor non-existent (Ehrendorfer.Whileit is conceivable generawith Andean-centered taxa.bothworking is in their groups. comm. forest inPleistocene and/or survival speciation from as resulting beeninterpreted Amazonian-centered entirely woody Prance(1973)selected For example. Andeanofspecific are documented by monographs endishia.Generaare typically that spemicrogeographic over 100species). vertically mountains.

librium (Richards.andKeister .Moreflower-feeding speciesin each Palaeotropical to 100-150 compared birdsare generally muchmorespecialized flower-feeding over.Madison. thePalaeotropics inepiphytes than richer aremuch The Neotropics has been to explain tendency the 1980). (Dodson. bird-flower it is probablethat humCertainly has been muchmorerapidin theNeotropics.'ca. tion. see Stiles.table5. alongthebase and lowerslopesofthenorthern concentrated speciation. 1975).Orchidaceae plant-pollinator 1980)seems pers. 69 are a common pollinators Shifts in specific thanthe exception. (1983)have suggested Andes. recentverydynamic Africa (and to a lesser Ratherthanthefloraof tropical extinction hypothesis.therather generthedata forflower-visiting Although to shownby Koopman(1981).compared (Land. forecologicaland altitudinal groups hummingbird-pollinated largely exactly as particular.97 in Venezuela. 1973.Terborgh side of the on thewestern concentrated is strikingly local endemism in general.1977. 1975).1981. phenomenally uniquely and as be considered rapid evidencethatthiskindof unusually circumstantial Thereis increasing theconin theNew Worldthanelsewhere. distributional patMexico.are clearlysimilar alized distributional patterns birdsand myAndean-centered planttaxa. a longhistory of mesicconditions from groupsresultsmostly Andean-centered and otherNorthern in epiphytes lack-ofof theprevalent almostthe antithesis speciation. earlier beganrelatively instead that inthis paper. withrespect extent Asia) beingimpoverished enriched. and Central Andeanregion ofthe suchpatterns It is notyetclearwhether are morea cause or an effect Anofthenorthern ofplanttaxa characteristic evolutionary explosion apparent and Winters (1982) have shownthatforbirds dean region. 1982).the Neotropical thantheir Palaeotropical equivalents.Thus Stiles(1981)has shownthatthereare over400 species of flower315 speciesofhummingbirds alone. 1970). in tropical partsof the northern and premontane are concentrated mingbirds Andeanregion(134 species in Colombia. 133 in Ecuador (Bleiweiss. terns) ofplants.However. a major evolutionary equihint ofanykindofecological theslightest without phenomenon open-ended or limits on birdsin theNeotropics. (at least to Africa) in the relatively thisdifference as due to lack of extinction as reflecting epiphyte diversity high and to interpret constantly mesicNeotropics I 1980).60 in 1978). proposethathighdiversity (Burger. wouldappearan altogether speciation In suchgroups theme. Ericaceae(Luteyn. bats are less precise. coevolution. including feeding realm..I wouldsuggest discussed patterns textofthebotanical speciaand hummingbird in general. thelatter may to the Neotropics. to and has not been restricted interaction has involvedmuchco-evolutionary plants.)orFuchsia(Berry.pers.). flower specificity and show muchgreater from thesepatterns that bird-flower coevolution probably Stiles(1981)suggested However.118 in Peru (Parkeret al.588 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. The greatest thoseof flower-visiting bat species(13 specieseach) are in thenorthern ofnectar-feeding concentrations America. However. reasonswhy theoretical northern et al.51 in Costa Rica. the rule rather ofprecise biotictuning rise to finer giving and co-evolution modeof speciation systems such as thosein Heliconia (Stiles. 37 in Guatemala Panama(Ridgely. to 52 in comm.

scale might on a smaller patterns similar although in thePalaeotropics. reflect burstof in Neogenetime.especially manyof the Neotropical along the proto-Antilles. is missing equivalent be expectedin New Guinea. phenomena The evolutionary evolution.whichseemstheclosestPalaeotropical oftheAndeancordilleras.mostly Uplift A similar explosion evolutionary families.led to an incredible of the Andes. givenriseto many flodirect forrelatively was a possibility there At theend of theCretaceous via island North America and tropical SouthAmerica between ristic interchange groups.and consistmostly withhumrelationships coevolutionary that suggest systems theirpollination Euglossines. (1982)notea grossly Marshall Moreover.are poorlyrepreAndeanregionand southern the northern and palmettos. that flora halfoftheNeotropical thisapproximately It is essentially Neotropics. miin thePlioceneled to (1) southward isthmus Closingof thePanamanian have becomeecologof someLaurasiantaxa intotheAndeswherethey gration at leastin woodytaxa. hopping may in bothregions. opwiththeconcomitant of theAndeanuplift. forexploiting potential ingtheevolutionary ofthe explainthe"excess" plantspeciesdiversity maylargely shrub strategies. seems that evolution evenexplosive dynamic. of theimmigrant versification here sugratesof speciation sociatedwiththeAndes.1982] GENTRY-NEOTROPICAL FLORISTIC DIVERSITY 589 should region in thisgeographical prevalent structures population thepartitioned similar et al. epiphytic. nectar-feeding mingbirds. Thus thehistorical plantshavtaxa ofGondwanan amongcertain forexplosive speciation portunity and understory palmetto. differentiation some of thedryarea taxa thatshowstrong thisearlyinterchange.The taxonomic in centers explosionhave distributional thisevolutionary thathave undergone CentralAmerica. of epiphytes. plantspecies appearto belongto thesegroups in thePalaeotroprepresented are muchmorepoorly all ofwhich andpalmettos. and speciation. sentedin Amazonia. little undergoing despite icallydominant . rapidcoevolution. favor inwhat landbridge oftheIsthmian resulting with formation formammals pattern faunasbetween of equilibrium to have been a balancedexchange theyconsider disecondary by a uniqueand unbalanced followed Northand SouthAmerica somehowasapparently taxa in SouthAmerica. theexceedingly To summarize. shrubs. taxa has givenriseto a verysigthenorthern-Andean-centered to characterize halfof all Neotropical flora. role in their have played a prominent half ofthetotal Neotropical account foralmost theAndeanuplift associatedwith richness oftheNeofortheexcess floristic and are thuslargely responsible flora tropics.Almost ofthetotalNeotropical proportion nificant shrubs. accident ics. Perhapsthe accelerated plantsis partof a muchmoregeneral SouthAmerican gestedfornorthwestern phenomenon. ofGondwanan in a number speciation groups inthesametaxaalso tookplace inCostaRica andPanama. as bees specialized such and perhaps bats. understory of epiphytes. CONCLUSION evolvedduring intherestofthetropics similar to that flora A richangiosperm has subsequently butthisflora in SouthAmerica thelast halfoftheCretaceous morespeciesin theNeotropics.

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