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The Effect of Nectar-Thieving Ants on the Reproductive Success of Frasera Speciosa (Gentianaceae)" (1988)

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The Effect of Nectar-Thieving Ants on the Reproductive Success of Frasera Speciosa (Gentianaceae) The College at Brockport: State University of New York Digital Commons @Brockport Environmental Science and Ecology Faculty Environmental Science and Ecology Publications 10-1988 The ffecE t of Nectar-Thieving Ants on the Reproductive Success of Frasera speciosa (Gentianaceae) Christopher J. Norment The College at Brockport, [email protected] Follow this and additional works at: https://digitalcommons.brockport.edu/env_facpub Part of the Environmental Sciences Commons Repository Citation Norment, Christopher J., "The Effect of Nectar-Thieving Ants on the Reproductive Success of Frasera speciosa (Gentianaceae)" (1988). Environmental Science and Ecology Faculty Publications. 24. https://digitalcommons.brockport.edu/env_facpub/24 Citation/Publisher Attribution: American Midland Naturalist, Vol. 120, No. 2 (Oct., 1988), pp. 331-336 This Article is brought to you for free and open access by the Environmental Science and Ecology at Digital Commons @Brockport. It has been accepted for inclusion in Environmental Science and Ecology Faculty Publications by an authorized administrator of Digital Commons @Brockport. For more information, please contact [email protected]. The University of Notre Dame The Effect of Nectar-Thieving Ants on the Reproductive Success of Frasera speciosa (Gentianaceae) Author(s): Christopher J. Norment Reviewed work(s): Source: American Midland Naturalist, Vol. 120, No. 2 (Oct., 1988), pp. 331-336 Published by: The University of Notre Dame Stable URL: http://www.jstor.org/stable/2426005 . Accessed: 10/04/2012 13:57 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]. The University of Notre Dame is collaborating with JSTOR to digitize, preserve and extend access to American Midland Naturalist. http://www.jstor.org The Effectof Nectar-thieving Ants on the Reproductive Success of Frasera speciosa (Gentianaceae) CHRISTOPHER J. NORMENT WestoverSchool, Middlebury, Connecticut 06762 ABSTRACT.-The impact of nectar-thievingants on the reproductivesuccess of Frasera speciosa (Gentianaceae), a perennial monocarp with periodic,synchronous flowering, was studied in a Wyoming alpine meadow. Plants fromwhich ants were excluded had higher rates of flowervisitation by other insects,and higher standing crops of nectar, than did plants visitedby ants. However, therewere no significantdifferences in eitherseed set or seed predationin plants with and withoutants. The lack of effectof ants upon the repro- ductivesuccess of Frasera may be due in part to temporalseparation of the activitiesof ants and some importantpollinators and seed predators.However, the abundant nectarproduc- tion, large inflorescencesand low rates of seed predationin Frasera,which are all related to its habit of periodicsynchronous flowering, may reduce the effectsof nectar-thievingants upon the species. INTRODUCTION Ants that feed on floral nectar pollinate several temperate plant species, including Po- lygonum cascadense (Hickman, 1974), Oreoxis alpina, Eritrichium aretioides and Thlaspi alpestre (Petersen, 1977), and Diamorpha smallii (Wyatt, 1981). However, ants may also act as nectar robbers or thieves (McDade and Kinsman, 1980; Willmer and Corbet, 1981; Schaffer et al., 1983), thus decreasing floral rewards without contributing to pollination. Nectar-robbing or -thieving ants (sensu Inouye, 1980) may reduce the reproductive success of the plants they visit. Inflorescences of Asclepias curassavica from which ants were excluded showed higher rates of pollinia insertion than did inflorescences visited by ants (Wyatt, 1980). Umbels and stems of Asclepias syriaca visited by ants initiated fewer pods than did umbels and stems without ants (Fritz and Morse, 1981). In a study of Pavonia dosypetala, flowers robbed by both bees and ants produced fewer seeds than flowers visited only by pollinating hummingbirds (Roubik, 1982). Galen (1983) found that nectar-thieving ants decreased seed production in Polemonium viscosum,a perennial alpine herb of western North America, by damaging the gynoecium during nectar foraging. Ants may also decrease reproductive success by impairing pollen function (Beattie et al., 1984). With few exceptions (Fritz and Morse, 1981; Fowler and Whitford, 1982; Galen, 1983), most studies of nectar- robbing and -thieving ants have occurred in tropical regions. In this article I examine the impact of nectar-thieving ants upon the reproductive success of monument plant (Frasera speciosa) (Gentianaceae), a long-lived perennial monocarp that grows in montane regions throughout western North America (Beattie et al., 1973). Specifically, I: (1) quantify ant visitation to Frasera flowers; (2) measure the impact of ants upon pollinator visitation and nectar resources and (3) examine the effectof ants upon seed set and seed predation. Frasera speciosa exhibits a pattern of synchronized flowering in which large amounts of elongated flowering stalks up to 3 m high are produced at 2-4-yr intervals (Beattie et al., 1973; Taylor and Inouye, 1985). Each inflorescence contains hundreds of campanulate- rotate, 2-5 cm broad, flowers. A large nectary, consisting of two canals up to 1 cm long, with fimbriate margins, is located at the base of each corolla lobe. Frasera flowers attract a 331 332 THE AMERICAN MIDLAND NATURALIST 120(2) large varietyof pollen- and nectar-feedinginsects, including coleopterans, dipterans, hy- menopteransand lepidopterans(Beattie et al., 1973). METHODS Ant-Frasera interactionswere studied between 1 July-20 August 1984 in an alpine meadow (elevation3050 m) at Clay Buttein the BeartoothMountains, Wyoming. In 1984, Fraserainflorescences were veryabundant, and plants floweredbetween 4 July-15 August. Twelve pairs of plants were selectedfor intensiveobservations; ants were excluded from one memberof each pair by coating the base with TanglefootG,tying up enough rosette leaves to preventant access (ca. fourout ofa minimumof 30 leaves) and clippingsurrounding vegetation.Since Frasera exhibitsdensity-dependent seed set (Taylor and Inouye, 1985), plants were selectedso that membersof each pair were within2 m of one another. At 3-d intervals,5-min censuses of flowervisitors were conductedon each plant. Im- mediatelyprior to observingeach plant,I countedall ants on the inflorescence.During each census the number and type of all flowervisitors except ants were recorded.Subsequent identificationof specimensfrom a referencecollection of Frasera visitors allowed mostinsects to be identifiedto the genus or species level (Table 1), althoughcommon dipterans could not be identifiedbeyond the order level while in the field.In addition to countingflower visitors,I also tallied all ant-flowervisitor interactions in which an ant caused a foraging insectto leave a flower. Frasera plants in the studypopulation floweredfrom 4 July-15 August,but to simplify comparisonsbetween treatments and individuals,data on visitationwere gatheredduring the 3-wk period when all stalks containedopen flowers.Each plant was censused seven times; total observationtime for each treatmentwas 420 min. To determineseed set and seed predation,50 matureovaries were collectedrandomly from all portionsof each plant, preservedin 70% ethanol, and later dissected.Values for the proportionof seed set and fruitsdamaged per plant were arcsin-transformedto correctfor nonnormalitybefore sta- tisticalanalysis. An exclusion experimentwas conductedto determinethe effectsof ants upon standing cropsof nectar.On each of foursampling dates, all flowervisitors were excludedfrom three plants near thoseused in the visitationexperiment by coatingthe base of the floweringstalk with TanglefootGand enclosingthe entireinflorescence in a fine-meshednylon bag. At the same time,the stalks of threeadjacent plants were coveredwith the same material staked to the ground; this preventedvisits by flyinginsects, but allowed free access by crawling insects. After24 h, six flowerswere collected from each inflorescenceand their nectar volumes measured to the nearest 0.1 ,l with a microsyringe.Nectar concentrations(total dissolvedsolids) were measured with a hand refractometerimmediately after collection of samples fromflowers with total accumulationsof >10 Al. RESULTS The mostcommon visitors to bothant-access and ant-exclusionplants were small dipterans (primarilyMuscidae and Bibionidae), a small nectar-gatheringbee (Evylaeus synthridis) and a nectar-feedingelaterid beetle (Ctenicerafallax) (Table 1). Workersof two bumblebee species (Bombus centralis and B. frigidus) were observed occasionally; a noctuid moth (Euxoa lewisi) was foundon plants in both treatments,although it was not seen duringthe timed censuses.An average of 34.4 ? 27.8 (SD) ants/plant,primarily Formica neorufibarbis, were counted on ant-accessplants. These ants were active mainly around floral nectariesand rarelycontacted stigmas or anthers.Heights and numberof flowersper inflorescencedid not differsignificantly between ant-access and ant-exclusionplants (Table 2). 1988 NORMENT: NECTAR-THIEVING ANTS 333 TABLE 1.-Flower visitorsto ant-access and ant-exclusionFrasera speciosa plants during timed censuses
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