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Bat/Hummingbird Pollination of a Cactus

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Bat/Hummingbird Pollination of a Cactus Bat and Hummingbird Pollination of an Autotetraploid Columnar Cactus, Weberbauerocereus weberbaueri (Cactaceae) Author(s): Catherine T. Sahley Source: American Journal of Botany, Vol. 83, No. 10 (Oct., 1996), pp. 1329-1336 Published by: Botanical Society of America Stable URL: http://www.jstor.org/stable/2446118 . Accessed: 03/12/2014 11:09 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]. Botanical Society of America is collaborating with JSTOR to digitize, preserve and extend access to American Journal of Botany. http://www.jstor.org This content downloaded from 192.135.179.249 on Wed, 3 Dec 2014 11:09:35 AM All use subject to JSTOR Terms and Conditions AmericanJournal of Botany 83(10): 1329-1336. 1996. BAT AND HUMMINGBIRD POLLINATION OF AN AUTOTETRAPLOID COLUMNAR CACTUS, WEBERBA UEROCEREUS WEBERBA UERI (CACTACEAE)' CATHERINE T. SAHLEY2 Departmentof Biology, Universityof Miami, Coral Gables, Florida 33124 I presentdata on flowermorphology, pollination, breeding system, and genetic diversityof the columnarcactus Weber- bauerocereus weberbaueriat 2500m elevation in southwesternPeru. Weberbauerocereusweberbaueri is a self-compatible columnarcactus thatis visited and pollinatedby one species of rare endemic bat, Platalina genovensium,and two species of hummingbirds,Patagona gigas and Rhodopis vesper. W. weberbaueriexhibits pronounced interplant variation in floral color and size, and flowersexhibit traitscorresponding to both bat and hummingbirdpollination syndromes.Starch-gel electrophoresisof flowerbud tissue indicated that W. weberbaueri is an autotetraploidand that genetic diversity(Hep = 0.257) of the studypopulation was high relativeto diploid plantsbut similarto othertetraploid species. Initial fruitset from pollinatorexclusion experimentsconducted in 1991 and 1993, at the onset of and aftera droughtassociated with the El Ninioevent of 1991-1992, revealed thatbats were the most importantpollinators in 1991, but thathummingbirds and diurnal insects were most importantin 1993. In both years,however, autogamy and lepidopteranlarval infestationof fruitsreduced differencesin maturefruit production among pollinatorexclusion treatmentsso thatdifferences in maturefruit set were not statisticallysignificant. Reduced bat pollinationin 1993 is attributedto the reduced abundance of bats at the study site duringa droughtcaused by El Niiio. I hypothesizethat interaction among several factors,including tetraploidy, autogamy, larval infestationof developing fruits,and variationin pollinatorabundance, may not resultin strongselection for a bat vs. hummingbirdfloral morph, thus allowing the persistenceof floralvariation in this cactus. Key words: Andean habitats;bats; Cactaceae; hummingbirds;Peru; pollination;polyploidy; Weberbauerocereus. In arid habitats,a large proportionof plantsbelonging share several characteristics,evolutionary transitions be- to the Cactaceae and Agavaceae exhibitfloral adaptations tween these syndromeswithin a plant lineage may occur for bat and/orhummingbird pollination (Porsche, 1939; relativelyfrequently (Helverson, 1993). In fact, several Vogel, 1969). Several recentfield studies in Mexico, Cu- plantfamilies (including the Cactaceae), genera,and even ragao, and Venezuela have experimentallydemonstrated species, exhibitfloral traits corresponding to bothbat and the importanceof bats (Nassar, 1991; Petit, 1995; Flem- hummingbirdpollination syndromes(e.g., Barthlottand ing, Tuttle,and Horner,in press) and/orhummingbirds Hunt, 1993; Helverson, 1993; Kress and Stone, 1993; (Fleming, Tuttle,and Horner,in press) as pollinatorsof Sazima, Sazima, and Buzato, 1994) yet few studieshave columnar cacti. Traits associated with bat pollinationof compared the relativecontribution of these pollinatorsto cacti include large, funnelform,nocturnal, white, and fruitand seed set. pungent-smellingflowers that produce large amountsof The purpose of this study was to investigatethe im- pollen and nectar."Hummingbird" flowers, on the other portanceof bat and hummingbirdpollination in Weber- hand, tend to be tubular,red, diurnal,and odorless, but bauerocereus weberbaueri (tribe Trichocereeae), a co- may also produce copious nectar and large amounts of lumnarcactus occurringin southwesternPeru thatexhib- pollen (Faegri and van der Pijl, 1979; Helverson, 1993). its floraltraits associated withboth bat and hummingbird Perhaps because bat and hummingbirdfloral syndromes pollination.Several factorsmotivated this study:first, the arid regions of Peru contain the second richest cactus floraafter Mexico, with -44 genera and 249 species dis- i Manuscript received 28 June 1995; revision accepted 21 March tributedprimarily along the westernAndean slopes (We- 1996. diversity, The authorthanks M. Donnelly, T. Fleming, J. Hamrick,D. Janos, berbauer,1945; Hunt, 1992). Yet, despite the C. Horvitz, K. Lips, M. Nelson, K. Waddington,and membersof the abundance, and prominence of these plants, virtually plant-animal interactionsgraduate discussion group of the University nothingis known regardingtheir ecology and reproduc- of Miami for valuable discussion, assistance, and comments;L. Bar- tive biology. Second, habitatsoccurring along thewestern aybar forassistance in the fieldthroughout the durationof the study;J. slopes of the Andes are underrepresentedwithin Peru's Estefaneroand J. Ugarte for assistance at the outset of the study; P nationalreserve system,and have been rankedas deserv- Jimenez for importantlogistic advice; E. Linares for assistance with researchand conservation(Centro plant identification;P Baldizan for graciouslypermitting the authorto ing of high priorityfor conduct researchon his property,and J. Hamrick of the Universityof de Datos para la Conservacion, 1986). Third, the only Georgia for kindlyallowing me the opportunityto work in his labora- nectarivorousbat occurringat mid-to-highelevation sites tory.Funds for this researchwere provided by grantsfrom New York in southwesternPeru, Platalina genovensium,is an ex- Zoological Society-TheWildlife Conservation Society, a WorldWildlife tremelyrare and endemic bat for which no ecological Fund-Garden Club of America Scholarship in Tropical Botany, Bat informationexisted prior to this study.Finally, W. we- ConservationInternational, and a Maytag GraduateFellowship fromthe varia- College of Arts and Sciences, Universityof Miami. berbaueri exhibitsconsiderable within-population 2 Currentaddress: CONATURA, Apartado 688, Arequipa, Peru (Fax: tion in several floralmorphological characters including 5154 424372). size, shape, and color, althoughthis variationhad never 1329 This content downloaded from 192.135.179.249 on Wed, 3 Dec 2014 11:09:35 AM All use subject to JSTOR Terms and Conditions 1330 AMERICAN JOURNAL OF BOTANY [Vol. 83 350 300 250 E 200 4- C 150 100 50 0n 62 63 64 65 66 67 68 69 70 71 81 82 83 84 85 86 87 88 89 90 91 92 93 Year Fig. 1. Rainfall data for 1962-1971 and 1981-1993 forthe Characato weatherstation (2400 m above sea level). This stationis 15 km from the main studysite and occurs withinthe same life zone. El Nifnoevents occurredin 1963-1965, 1969, 1972-1973, 1976, 1982-1983, and 1991- 1992 (no rainfalldata was available for 1969 and 1972-1973 events). Rainfall data from1961 to 1976 fromONERN, 1974. previouslybeen quantified.Among-plant variation in flo- tus in the region around the city of Arequipa (Aragon, 1982; C. T ral color ranges frombright pink-red to white.This vari- Sahley, personal observation).Although the populationaround Arequi- ation in floral color, along with the presence of long, pa has been classifiedas threespecies by Ritter(1981), the continuous stout,funnelform-tubular corollas and both diurnal and variationamong plants in flowermorphology and color and the lack of nocturnalnectar production, suggests adaptationto both any diagnosticalleles among morphs,has led me to consider them as bat and hummingbirdpollination. Prior to this study, one highlyvariable species, W. weberbaueri,in accordance with Hunt however,the relativeimportance of bat and hummingbird (1992). pollination for fruitproduction in W. weberbaueri was Flowers of W. weberbaueri are hermaphroditic,begin opening be- unknown. tween 1500 and 1700, remainopen throughoutthe night,and generally To elucidate possible ecological and genetic mecha- close by 1100 the followingmorning. Nectar productionis continuous, but variable, while the floweris flower nisms contributingto the presence of floralmorphologi- open (Sahley, 1995). Although and fruitproduction peaks in Septemberand October,some flowerand cal variationin this cactus, I addressedthe followingfive fruitproduction occurs year-round(Sahley, 1995). Weberbauerocereus questions: (1) What is the morphologicaland geneticdi- weberbaueriis visitedby the rare and endemic nectar-feedingbat, Pla- versityof W. weberbaueri at the studysite? (2) What is talina genovensium(Phyllostomidae), two species of hummingbirds, the extentof outcrossingvs. selfingoccurring within the Patagona gigas and Rhodopis vesper (Trochilidae),small, unidentified studypopulation? (3) What is the relativeimportance of nocturnalmoths,
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