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The Ant, Aphaenogaster Picea, Benefits from Plant Elaiosomes When Insect Prey Is Scarce Author(S): Robert E

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The Ant, Aphaenogaster Picea, Benefits from Plant Elaiosomes When Insect Prey Is Scarce Author(S): Robert E The Ant, Aphaenogaster picea, Benefits from Plant Elaiosomes When Insect Prey is Scarce Author(s): Robert E. Clark and Joshua R. King Source: Environmental Entomology, 41(6):1405-1408. 2012. Published By: Entomological Society of America URL: http://www.bioone.org/doi/full/10.1603/EN12131 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/ terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. PLANT-INSECT INTERACTIONS The Ant, Aphaenogaster picea,BenefitsFromPlantElaiosomesWhen Insect Prey is Scarce 1,2 3 ROBERT E. CLARK AND JOSHUA R. KING Environ. Entomol. 41(6): 1405Ð1408 (2012); DOI: http://dx.doi.org/10.1603/EN12131 ABSTRACT Myrmecochory is a facultative, mutualistic interaction in which ants receive a protein- rich food reward (elaiosome) in return for dispersing plant seeds. In North American northeastern hardwood forests, Aphaenogaster ants are the primary genus dispersing myrmecochorous plants. In these forests, myrmecochores occur in plant guilds of understory spring ephemerals or seasonal greens. This mutualism has been demonstrated for Aphaenogaster rudis (Emery) and individual plant species, but it has not been demonstrated for other Aphaenogaster species or guilds of myrmecochores as they naturally occur. Aphaenogaster picea (Wheeler) colonies were fed three treatments over 5 mo: 1) a mixture of only elaiosomes from an entire plant guild, 2) a diet of only insect protein and 3) a combination diet of both elaiosomes and insect protein. This experiment investigated two potential hypotheses through which elaiosomes can beneÞt ants: 1) elaiosome proteins can substitute for protein nutritional requirements when ants are prey-limited, and 2) elaiosome nutrition can supplement insect protein when prey is ample. First, a mixture of elaiosomes from four myrmecochorous plant species provided to A. picea colonies was sufÞcient to maintain worker production, larval growth, and fat stores when no other food was available. A. picea colonies consuming elaiosomes as their only protein source could be sustained for a growing season (5 mo). Second, colonies fed both elaiosomes and protein did not yield more productive colonies than a control diet of just insect protein. These results support the hypothesis that myrmecochory is indeed a facultative mutualism in which ants take advantage of the protein content of elaiosomes when it is favorable, but when they are not limited by insect prey they do not gain any additional beneÞt from elaiosomes. KEY WORDS antÐplant interactions, myrmecochory, elaiosome, mutualism, trophic interactions Myrmecochory is a mutualism where ants disperse diet of the ant mutualist: in addition to scavenging plant seeds in return for a protein-rich reward called insect protein and elaiosomes, it has been shown that an elaiosome (Rico-Gray and Oliveira 2007). In North eastern Aphaenogaster species take insect prey as an American northeastern hardwood forests Aphaeno- important food source (unpublished data, Buczkowski gaster ants are the primary dispersers of myrmeco- and Bennett 2007). chorous plants (Ness et al. 2009). Generally speaking, The Aphaenogaster fulvaÐrudisÐtexana species com- the beneÞt of myrmecochorous seed dispersal is con- plex (Umphrey 1996) is a group of abundant and text speciÞc: the plant may gain a nutrient rich mi- widespread ants in North American northeastern for- crosite (Beattie and Culver 1982) or protection from ests. For this study, in Connecticut, U.S.A., the species predation (Heithaus 1981), competition (Handel Aphaenogaster picea (Buckley) is an abundant mem- 1976), or Þre (Rico-Gray and Oliveira 2007). ber of this species complex. Like other members of the Aphaenogaster rudis ants are shown to beneÞt from Aphaenogaster fulvaÐrudisÐtexana complex, A. picea this association with myrmecochores (Heithaus et al. disperses myrmecochorous plant seeds (unpublished 2005). This is because elaiosomes are an important data). A. picea is a widespread ant of mesic deciduous food source for Aphaenogaster rudis colonies (Morales forests in eastern North America. Umphrey (1996) and Heithaus 1998, Bono and Heithaus 2002, Fischer described the range of A. picea, which typically is et al. 2008, Fokuhl et al. 2007), providing protein and associated with inland, relatively higher elevations some lipid content. Like other facultative insect mu- than the Atlantic coast and spans the range from Con- tualisms, this food reward does not make up the entire necticut and Ontario, Canada; west to Pennsylvania, south into the southeastern United States along the 1 Wesleyan University, Department of Biology, Middletown, CT Appalachian Mountains. To date no study has shown 06457. that members of this species complex, other than 2 Corresponding author: Robert E. Clark, Wesleyan University, Aphaenogaster rudis, beneÞt from this mutualism. Biology Department, 52 Lawn Ave., Middletown, CT 06457 (e-mail: A. picea co-occurs with several myrmecochorous [email protected]). 3 University of Central Florida, Biology Department, Orlando, FL, species (ranges published by Umphrey 1996 and 32816. Kartesz 2011). Multiple species of myrmecochores 0046-225X/12/1405Ð1408$04.00/0 ᭧ 2012 Entomological Society of America 1406 ENVIRONMENTAL ENTOMOLOGY Vol. 41, no. 6 overlap within the same habitats at small spatial scales and make up an appreciable diversity and biomass of understory herbs (Handel et al. 1981, unpublished data, R.C.). Because of the frequent high density of both Aphaenogaster and myrmecochores and poten- tially spatially overlapping seed shadows, ant colonies may consume multiple plant species elaiosomes over the course of a season. For this reason, feeding exper- iments that use one plantÕs elaiosomes do not reßect the food rewards typically experienced by one ant colony. In these forests myrmecochory is a species- guild mutualism not a species-species mutualism. Therefore to determine the nutritional beneÞts of myrmecochory for ants it is realistic to provide a mix- ture of elaiosomes. In this experiment the mixtures chosen reßect the guild of myrmecochores A. picea interacts with. Does a guild of myrmecochores provide a tangible beneÞt to ant colonies? We tested whether or not a mixture of elaiosomes provide enough nutrition to have a signiÞcant impact on worker production (by Fig. 1. Distribution of colony sizes in each experimental observing brood numbers and worker count) and en- treatment. Total adult worker count in each A. picea colony ergy (by observing lipid stores). This feeding treat- varied from 15 to 450 workers. Colonies in the same treat- ment was achieved under conditions with strict diet ment have connected bars. Ant colonies were stratiÞed ran- control: testing for the beneÞt of elaiosomes when domly between three treatments based on size. insect protein was limiting and the beneÞts of elaio- somes when insect protein was ample. A. picea colo- nies across different feeding treatments were reared Cassill 2012). Colonies were cooled and then kept in under laboratory conditions. atemperatureregulatedtoensurethiscontrol.Acool- ing period, however, accounts for potential cold tem- perature triggers normally experienced by ants en- Materials and Methods demic to temperate regions. Before experimental From May to September of 2009, A. picea colonies treatments in December 2009, A. picea colonies were were collected from secondary-growth, mesic decid- chilled in a 2ЊClabrefrigerator.FortheÞrstweek, uous forests of Connecticut, U.S.A. Colony rearing colonies were removed from the refrigerator every 2 d methods follow Lubertazzi and Adams 2010, who stud- for 12 h. Colonies were not fed during this cooling ied the role of nutrition in Aphaenogaster rudis colony cycle to mimic the natural nutritional limitations of growth and sex allocation. In this way, the ants were winter. cultured into ßuon-lined plastic boxes with moist den- All seeds, with elaiosomes, were gathered opportu- tal plaster bottoms, and provided water, dead crickets, nistically as seed pods started to dehisce from May to and 20% sucrose, ad libitum. Temperature was main- August of 2009. These were collected from the same tained at a constant 30ЊC. Variation in temperature can local Connecticut forests where colonies were col- confound colony growth responses to food sources lected. The feeding treatments represent a distribu- (Asano and Cassill 2012). Twenty complete colonies tion of elaiosomes that would typically occur with A. (with queens) were maintained under these condi- picea and included the four most common mymrem- tions until the experimental treatments were started in cochores in these habitats (unpublished data, R.C.). December 2009. For each given colony, sexual castes These elaiosome-bearing
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