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Consequences of the Timing of Seed Release of Erythronium Americanum (Liliaceae), a Deciduous Forest Myrmecochore Author(S): Scott Ruhren and Michele R

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Consequences of the Timing of Seed Release of Erythronium Americanum (Liliaceae), a Deciduous Forest Myrmecochore Author(S): Scott Ruhren and Michele R Consequences of the Timing of Seed Release of Erythronium americanum (Liliaceae), a Deciduous Forest Myrmecochore Author(s): Scott Ruhren and Michele R. Dudash Source: American Journal of Botany, Vol. 83, No. 5 (May, 1996), pp. 633-640 Published by: Botanical Society of America Stable URL: http://www.jstor.org/stable/2445923 Accessed: 10/06/2010 10:37 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=botsam. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. 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 AmericanJournal of Botany 83(5): 633-640. 1996. CONSEQUENCES OF THE TIMING OF SEED RELEASE OF ERYTHRONIUMAMERICANUM (LILIACEAE), A DECIDUOUS FOREST MYRMECOCHORE' SCOTT RUHREN2,3 AND MICHELE R. DUDASH Departmentof Botany,University of Maryland,College Park, Maryland 20742 Myrmecochoresare plants with seeds adapted for ant dispersal. This specialized dispersal syndromemay provide Ery- throniumamericanum seeds withprotection from predators within the easterndeciduous forests.To determinethe adaptive significanceof myrmecochoryin E. americanum,seed removal rates and seed predationin relationto seed release date and location along the Potomac River in Langley, Virginia,were examined. The numberof seeds removedfrom four exclosure treatmentswere monitoredtwo times in 1992 and three times in 1993 within floodplain and hilltop populations of E. americanum.Overall, seed removal was greatestfrom control depots, and E. americanumseeds were removedat nearlythe same rate frompredator-exclusion depots, indicatingthat removal fromopen depots is largely due to ant removal. Ants removed significantlymore seeds than predatorsin the first48 h of seed exposure and could potentiallyremove all E. americanumseeds before nightfall.Aphaenogaster rudis was identifiedas the primarydisperser of E. americanum.Seeds placed in depots afterthe naturalseed release period were discovered more quickly and removedby ants at a significantly higherrate than seeds released at the naturaldate. These resultssuggest that ant dispersal of E. americanumseeds reduces the likelihood of seed predation. Key words: Ant dispersal;deciduous forests;Erythronium americanum; Jeffersonia diphylla; Liliaceae; myrmecochores; seed release. Myrmecochory,the dispersal of seeds by ants, is a 1973; Oostermeijer,1989). Withinthe easterndeciduous widespread phenomenon documented in more than 70 forestsmany myrmecochoresrelease theirant-dispersed plant families (Beattie, 1985) and numeroushabitats in- seeds during a relativelyshort period. Some myrmeco- cluding deciduous forests (e.g., Holland, 1974; Culver chores exhibitoverlapping seed release phenologies dur- and Beattie, 1978; Muller, 1978, 1990), dunes (Ooster- ing thistime, with the less abundantseeds oftenpreferred meijer,1989), tropicalepiphyte communities (Kleinfeldt, by dispersersand predators(e.g., Smithet al., 1989). For 1978), and sclerophylous shrub (Bond and Slingsby, othermyrmecochores, however, there appears to be some 1984; Hughes and Westoby, 1990). Examples of myr- temporal separation of release dates (e.g., Beattie and mecochoryare most abundantin Australiaand South Af- Culver, 1981; Handel, Fisch, and Schatz, 1981; Heithaus, rica (Hughes and Westoby,1990; Westobyet al., 1991), 1986; Ruhren, 1994). For example, Erythroniumameri- but this syndromeis well representedin the easternde- canum Ker. (Gleason and Cronquist, 1991) is a typical ciduous forestsof NorthAmerica where myrmecochores ephemeralmyrmecochore exhibiting an elaiosome (Hol- may constituteup to 40% of the herbaceous layer (Han- land, 1974; Muller, 1978; Harder et al., 1985; Wein and del, Fisch, and Schatz, 1981). Pickett,1989) and a synchronousseed release period that Many of these myrmecochoresare co-occurringspring occurs afterViola spp. yet beforeseveral othersympatric ephemerals,herbaceous plants thatgrow rapidlyfollow- myrmecochoresin northernVirginia (e.g., Jeffersoniadi- ing snowmeltand begin to senesce as the canopy closes. phylla and Trillium sessile) (Lobstein and Rockwood, The prevalence of this syndromein the spring flora of 1993; Ruhren, 1994). This temporal separationof seed the easterndeciduous forestsmay be maintainedby the release among myrmecochoresmay maximize exposure paucity of lipid-richfoods (Carrol and Janzen, 1973). to seed-dispersingants. Therefore,seed-dispersing ants are attractedto the lipids It is theorizedthat myrmecochory may provide seeds contained in most elaiosomes (e.g., Carrol and Janzen with several benefits,including protection from seed I Manuscriptreceived 21 March 1995; revision accepted 2 October predators(e.g., Culver and Beattie, 1978; O'Dowd and 1995. Hay, 1980; Heithaus, 1981; Turnbulland Culver, 1983), The authorsthank C. DiSalvo and D. Sealy at USDOI, NPS for site avoidance of interspecificcompetition (e.g., Handel, access to TurkeyRun Park and cooperation;J. Wilkinsonand R. Iras- 1976, 1978), deposition in a nutrient-enrichedsafe site quin forfield assistance; taxonomistsat BeltsvilleAgricultural Research (Culver and Beattie, 1980, 1983) and protectionfrom fire Center,USDA for ant identityverification; D. Carr, C. B. Fenster,L. Galloway, S. N. Handel, E. Nagy, and K. C. Ruhrenfor commentson (e.g., Berg, 1975). Withinthe eastern deciduous forests the manuscript;and M. McIntosh, D. Denmann, D. Carr, and L. M. of North America, a drivingforce in this plant-animal Rosa for statisticaladvice. All analyses were run on the mainframeat interactionis seed predation.The elaiosomes thatattract the UMD ComputerScience Center.This studywas partiallyfunded by the ants also attractpredators including rodents, primarily a research award from the WashingtonBiologists' Field Club to S. Peromyscus leucopus (e.g., Heithaus, 1981), birds (Mi- Ruhren. 2 Currentaddress: Departmentof Biological Sciences, RutgersUni- lewski, 1982), and many arthropods(e.g., Bond and versity,Piscataway, New Jersey08855-1059. Slingsby, 1984; Anderson,1988). Most of the arthropod 3 Authorfor correspondence. seed predatorsare nocturnalground beetles (e.g., Ohara 633 634 AMERICAN JOURNAL OF BOTANY [Vol. 83 and Higashi, 1987; Higashi and Ito, 1991). Rodents are E. americanum -9 X 15 m mixed with Galium spp. and the myrme- assumed to be destructivebecause they do not cache cochores C. virginicaand A. canadense. seeds in the springand early summer(Barry, 1976). Hei- thaus (1981) discovered thatP. leucopus may cue in on Plant analysislseed collection-All ramets were counted and the the naturalseed depots formedunder dehiscing capsules floweringramets were marked with plastic stakes and monitoredto and frequentlydestroy predehiscent fruits. However, it determinedates of opening,onset of senescence, and cessation of fruit also was revealed that seeds are less detectable by P. and seed production.These individualswere checked in 1992 and 1993 leucopus afterants stripseeds of theirelaiosomes within to determinereproductive output. All capsules withinthe floodplainand theirnests. Thereforeseeds discovered and collected by hilltop were bagged in bridal veil, a fine mesh fabric,to preventpre- ants should exhibitreduced predation. dispersal predationand dispersal by ants and to facilitaterecovery of The goals of our study were to examine factors af- seeds. All seeds were counted and weighed. Seeds were pooled within fectingE. americanumseed dispersal and to evaluate the sites, and additional seeds were collected and weighed fromadjacent nonstudypopulations withinTRP. Seeds were refrigeratedfor several theoreticalbenefit of this dispersal mechanismin reduc- days to preventlipid degradationuntil sufficientseed was available to ing predationand its response to an artificiallyextended begin field manipulationsand observations.Seeds of this species that seed release phenology.First, we testedthis with a series had been refrigeratedfor 1 yr were detected and quickly removed by of treatmentsdesigned to exclude eitherants, rodents, or ants in 1993 (Ruhren,1994). both from seeds placed in two sites in both 1992 and 1993. Second, we extendedthe seed release period of E. Monitoringseed dispersal-In 1992 two transectswere placed 5 m americanumto evaluate the effectof timingon seed dis- apartin boththe floodplainand hilltopstudy sites to determineE. amer- persal and predationrates. Third,we compared removal icanum seed removal rates by ants and predators.The linear transects rates of E. americanum to Jeffersonia diphylla, a sym- contained seven replicates. Each replicate contained four exclusion
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