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Diversity and Biogeography of Neotropical Vascular Epiphytes Author(S): Alwyn H

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Diversity and Biogeography of Neotropical Vascular Epiphytes Author(S): Alwyn H Diversity and Biogeography of Neotropical Vascular Epiphytes Author(s): Alwyn H. Gentry and C. H. Dodson Reviewed work(s): Source: Annals of the Missouri Botanical Garden, Vol. 74, No. 2 (1987), pp. 205-233 Published by: Missouri Botanical Garden Press Stable URL: http://www.jstor.org/stable/2399395 . Accessed: 04/10/2012 16:06 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Missouri Botanical Garden Press is collaborating with JSTOR to digitize, preserve and extend access to Annals of the Missouri Botanical Garden. http://www.jstor.org DIVERSITY AND BIOGEOGRAPHY OF NEOTROPICAL VASCULAR EPIPHYTES' ALWYN H. GENTRY AND C. H. DODSON2 In his classic work Schimper (1888), empha- epiphyticranks. Only 32 seed plantfamilies have sizingthe taxonomicdiversity of epiphytes,list- as many as fiveor more epiphyticspecies, 26 of ed 33 familiesand 232 generaof epiphytes.Until these with epiphytesin the Neotropics. It is on veryrecently, subsequent authors have generally the 42 families(Table 3) that contain epiphytes accepted Schimper'sfigures (Richards, 1957; Jo- in the Neotropics that this paper will focus. hansson, 1974). However,epiphytism (here used Even thoughthis analysis of epiphytediversity in a broad sense to include hemiepiphytes,see and distributionis largelyfocused on the Neo- Kress, 1986) was recentlyreported to exist in 65 tropics,a fewcomparisons with the Paleotropics differentvascular plant families(56 familiesex- are instructive.There are actually slightlymore luding ferns),38 of these with epiphytesin the familieswith epiphytesin the Paleotropics (43) Neotropics(Madison, 1977). Our own data (Ap- than in the Neotropics (42), with all of the pa- pendix 1) and additional records compiled by leotropical epiphyticfamilies having epiphytic Kress (1986) now record83 vascular plant fam- representativesin Australasia but only 15 in Af- ilies withepiphytic species.3 At least 876 genera rica and Madagascar. If only the 32 seed plant include at least one epiphyticspecies and there familieswith five or more epiphyticspecies are are perhaps 29,000 epiphyticspecies, ca. 10% of considered,there are also roughlyequal repre- all vascular plants (Table 1). Thus at firstsight sentationsof epiphyte-containingfamilies in the epiphytismseems a very widespread and suc- Neotropics (26) and Australasia (25) but only cessfullife-style, which very many unrelated taxa about half as many in Africa(14). have evolved. At the species level the storyis verydifferent. However, a closer examination suggeststhat There are many more epiphytesin the Neotrop- even though there are both many species and ics, at least half again as many as in Australasia highertaxa of epiphytes,few of the highertaxa and six times as many as in Africa. Although account for most of the species. Burger(1985), similarnumbers of genera and familiesevolved forexample, emphasized thatrelatively few lin- epiphytismin the differentregions, subsequent eages have been able to enterthe epiphytic niche, speciationas epiphyteswas dramaticallygreater presumablybecause of the complex suite of ad- in the Neotropics. A major objective of this pa- aptations needed. Thus even though it is true per, then,will be to tryto explain why there is thatthe evolutionof an epiphytichabit has been so much epiphytediversity in the Neotropics. a relativelycommon featureof vascular plant evolution,it is equally true that very few of the EPIPHYTE FAMILIES taxa that have evolved an epiphytichabit have One approach to an overview of neotropical radiated successfullyto produce otherepiphytic epiphytediversity is a taxonomic one. Table 3 species (Table 2). In most of the epiphyte-con- summarizesthe neotropicalepiphytic seed plant tainingfamilies, epiphytism is a ratherinsignif- floraby family.Of the 42 families represented icant anomaly. Indeed, eliminatinga mere 85 by at least one habituallyepiphytic species in the such "oddball" species from the roster of the Neotropics,the Orchidaceae are by farthe most world's epiphytesremoves 31 familiesfrom the importantwith ten timesas manyepiphytic neo- I We thankthe National Science Foundation (INT-790-6840; BSR-834-2764) forsupport of our Ecuadorian fieldwork.Drs. H. Bedell, T. Croat, B. Hammel, J. Luteyn,S. Renner, P. Taylor, L. B. Smith, M. Dillon, and H. Robinson provided unpublished data on distributionof epiphytismin their taxonomic specialties. We especially thank B. Hammel formaking available unpublisheddata on the habit composition of the La Selva flora;B. Burger,T. Croat, B. Hammel, M. Madison, S. Renner, and E. Zardini for reviewingthe manuscript; and E. Zardini formuch technicalassistance in its preparation. 2 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. 3The data set used in this paper was derived independentlyfrom that of Kress (1986) and is mostlybased on Madison's figuresas modifiedin Appendix 1. ANN. MissouRI BOT. GARD. 74: 205-233. 1987. 206 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 74 TABLE 1. Taxonomic distributionof vascular epi- and suggested20,000-25,000 as the best esti- phytes(modified from Madison, 1977 and Kress, 1986). mate of orchid species numbers. Since 70% of the total number of orchid species should ap- Fami- Gen- proximate the number of species of epiphytic lies era with with Species orchid,there should be betweenca. 12,000 (from Epi- Epi- of Epi- AiryShaw's estimate)and ca. 20,000 (fromMad- phytes phytes phytes ison's estimate).Unfortunately, the 8,000 species Pteridophytes 13 92 2,593 "slop" betweenthese two estimatesis as greatas Gymnosperms 2 2 4 thetotal number of epiphytes in all otherfamilies combined!Madison (1977) used thehigher figure Monocots forhis calculations,whereas Dressler (1981) gen- Withoutorchids 80 2,657 erallyopted for more conservativeestimates of Orchids (fide Madi- son) 500 20,000 orchid species numbers.We have mostlytaken Orchids (compiled the highervalues since we know of many cases fromDressler, whereDressler's figuresfor species numbersare 1981) 460 15,000 significantunderestimates but none wherehe has Orchids (fide Kress, overestimated.For example, Dressler suggested 1986) 440 13,951 that there are 830 species in subtribe Epiden- Total (fideKress, drineae,whereas in Ecuador alone thereare 500 1986) 17 520 16,608 species in the single genus Epidendrum and we 17 540 Total (our estimate) 22,657' think 1,200 species would be a bettersubtribal Dicots 51 262 4,251 estimate. Total 83 896 29,505 Moreover, new orchid species are being dis- 1 Includes Madison's orchid figure(see text). covered at an astonishingrate, especially in the northernAndean region, again suggestingthat Dressler'sestimates of species numberswill have tropicalspecies as runners-upAraceae and Bro- to be adjusted dramaticallyupward. For exam- meliaceae. All threeof these most speciose neo- ple, about 2,315 orchid species are now known tropical epiphyte families are monocots. One fromEcuador, ca. 700 of these describedonly in other monocot family,Cyclanthaceae, also has the last 15 years.Nevertheless, more than 1,500 a significantnumber of epiphytes.Commelina- additional Ecuadorian orchid "morphospecies" ceae, Rapateaceae, and Philesiaceae, although have notbeen identifiedwith any published name. with few species, have an exclusivelyepiphytic Even if the 300 unaccounted fornames should genus (respectively,Cochilostemon, Epidryos, all prove applicable to the unidentifiedspeci- Luzuriaga, and (in our experience)Philesia). As mens at hand, it is inevitable that most of the summarizedby Madison (1977), the rest of the unidentifiedtaxa will prove undescribedand the epiphyticmonocot floraof the Neotropics con- list of orchids forEcuador alone will increase to sists of single species of Burmannia and Yucca well over 3,000 species. and a few Central American species of Smila- The neotropicalepiphytic dicot florais more cina. diversein familiesbut much less diversein species Since orchidsare so overwhelminglythe most than the monocots. Twenty-ninedicot families diverse group of epiphytes(about 70% of their have at least one habituallyepiphytic species in species are epiphytic),estimates of orchiddiver- the Neotropics. The largestof these are Pipera- sityare criticalto an evaluation of epiphytedi- ceae (ca. 500 spp.), Gesneriaceae (483 spp.), Me- versity. Unfortunately,orchids are amazingly lastomataceae(227 spp.), Ericaceae (ca. 300 spp.), poorly known taxonomically (compare the ca. Cactaceae (133 spp.), Guttiferae(ca. 90 spp.), 12 orchid taxonomistswith the ca. 200 system- and Marcgraviaceae (87 spp.). In addition there atists specializingin the similar-sizedCompos- are perhaps 110 neotropicalspecies of Moraceae itae). Estimatesof the numberof orchid species stranglersin the genera Ficus and Coussapoa. range from 12,000 ("some authors" fide Dress- The only otherdicot familieswith more than 20 ler, 1981) or 17,000 (AiryShaw, 1973) to 30,000 epiphyticspecies in the Neotropics are Aralia- (Madison, 1977) or 35,000 ("some authors" fide ceae, Bignoniaceae,Compositae, Rubiaceae, and Dressler, 1981). In the best available review Solanaceae (Table 3). Dressler (1981) counted almost
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