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Whether microbes also play a role in more modest host shifts or expansions in herbivorous insects is less clear. Here we evaluate the potential for gut microbial communities to constrain or facilitate host plant use in the Melissa blue butterfly (<jats:italic>Lycaeides melissa<\/jats:italic>). We conducted a larval rearing experiment where caterpillars from two populations were fed plant tissue from two hosts. We used 16S rRNA sequencing to quantify the relative effects of sample type (frass versus whole caterpillar), diet (plant species), butterfly population and development (caterpillar age) on the composition and diversity of the caterpillar gut microbial communities, and secondly, to test for a relationship between microbial community and larval performance. Gut microbial communities varied over time (that is, with caterpillar age) and differed between frass and whole caterpillar samples. Diet (host plant) and butterfly population had much more limited effects on microbial communities. We found no evidence that gut microbe community composition was associated with caterpillar weight, and thus, our results provide no support for the hypothesis that variation in microbial community affects performance in <jats:italic>L<\/jats:italic>. <jats:italic>melissa<\/jats:italic>.<\/jats:p>","DOI":"10.1038\/s41598-017-11781-1","type":"journal-article","created":{"date-parts":[[2017,9,6]],"date-time":"2017-09-06T22:13:41Z","timestamp":1504736021000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":30,"title":["Sources of Variation in the Gut Microbial Community of Lycaeides melissa Caterpillars"],"prefix":"10.1038","volume":"7","author":[{"given":"Samridhi","family":"Chaturvedi","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexandre","family":"Rego","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lauren K.","family":"Lucas","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zachariah","family":"Gompert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2017,9,12]]},"reference":[{"key":"11781_CR1","doi-asserted-by":"publisher","first-page":"1441","DOI":"10.1126\/science.241.4872.1441","volume":"241","author":"RM May","year":"1988","unstructured":"May, R. M. How many species are there on earth? Science \n                           241, 1441\u20131449 (1988).","journal-title":"Science"},{"key":"11781_CR2","unstructured":"Schoonhoven, L. M., van Loon, J. J. A. & Dicke, M. Insect-Plant Biology, second edn. (Oxford University Press, 2010)."},{"key":"11781_CR3","doi-asserted-by":"crossref","unstructured":"Jaenike, J. Host specialization in phytophagous insects. Annual Review of Ecology and Systematics 243\u2013273 (1990).","DOI":"10.1146\/annurev.es.21.110190.001331"},{"key":"11781_CR4","doi-asserted-by":"publisher","first-page":"442","DOI":"10.1073\/pnas.1423042112","volume":"112","author":"ML Forister","year":"2015","unstructured":"Forister, M. L. et al. The global distribution of diet breadth in insect herbivores. Proceedings of the National Academy of Sciences \n                           112, 442\u2013447 (2015).","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"11781_CR5","doi-asserted-by":"publisher","first-page":"555","DOI":"10.1126\/science.281.5376.555","volume":"281","author":"BD Farrell","year":"1998","unstructured":"Farrell, B. D. \u201cInordinate fondness\u201d explained: Why are there so many beetles? Science \n                           281, 555\u2013559 (1998).","journal-title":"Science"},{"key":"11781_CR6","doi-asserted-by":"publisher","first-page":"471","DOI":"10.1098\/rstb.2002.1059","volume":"357","author":"M Dr\u00e8s","year":"2002","unstructured":"Dr\u00e8s, M. & Mallet, J. Host races in plant\u2013feeding insects and their importance in sympatric speciation. 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Proceedings of the Royal Society of London B: Biological Sciences \n                           277, 3735\u20133743 (2010).","journal-title":"Proceedings of the Royal Society of London B: Biological Sciences"},{"key":"11781_CR9","doi-asserted-by":"publisher","first-page":"E5980","DOI":"10.1073\/pnas.1424717112","volume":"112","author":"GR Hood","year":"2015","unstructured":"Hood, G. R. et al. Sequential divergence and the multiplicative origin of community diversity. Proceedings of the National Academy of Sciences \n                           112, E5980\u2013E5989 (2015).","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"11781_CR10","doi-asserted-by":"publisher","first-page":"559","DOI":"10.1016\/S0965-1748(00)00024-2","volume":"30","author":"JL Stevens","year":"2000","unstructured":"Stevens, J. L., Snyder, M. J., Koener, J. F. & Feyereisen, R. Inducible P450s of the CYP9 family from larval Manduca sexta midgut. Insect Biochemistry and Molecular Biology \n                           30, 559\u2013568 (2000).","journal-title":"Insect Biochemistry and Molecular Biology"},{"key":"11781_CR11","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1046\/j.1365-2583.2003.00395.x","volume":"12","author":"K Zhu-Salzman","year":"2003","unstructured":"Zhu-Salzman, K., Koiwa, H., Salzman, R., Shade, R. & Ahn, J.-E. Cowpea bruchid Callosobruchus maculatus uses a three-component strategy to overcome a plant defensive cysteine protease inhibitor. Insect Molecular Biology \n                           12, 135\u2013145 (2003).","journal-title":"Insect Molecular Biology"},{"key":"11781_CR12","doi-asserted-by":"publisher","first-page":"298","DOI":"10.1016\/j.tree.2007.02.010","volume":"22","author":"L Despres","year":"2007","unstructured":"Despres, L., David, J.-P. & Gallet, C. The evolutionary ecology of insect resistance to plant chemicals. Trends in Ecology & Evolution \n                           22, 298\u2013307 (2007).","journal-title":"Trends in Ecology & Evolution"},{"key":"11781_CR13","doi-asserted-by":"publisher","first-page":"1989","DOI":"10.1126\/science.1094611","volume":"303","author":"T Tsuchida","year":"2004","unstructured":"Tsuchida, T., Koga, R. & Fukatsu, T. Host plant specialization governed by facultative symbiont. Science \n                           303, 1989\u20131989 (2004).","journal-title":"Science"},{"key":"11781_CR14","unstructured":"Berenbaum, M. Allelochemicals in insect-microbe-plant interactions: Agents provocateurs in the revolutionary arms race. Novel Aspects of Insect-Plant Interactions, eds Barbosa, P. & Letourneau, D. K. (Wiley, New York) 97\u2013123 (1988)."},{"key":"11781_CR15","doi-asserted-by":"publisher","first-page":"533","DOI":"10.1111\/j.1365-2311.2011.01318.x","volume":"36","author":"H Feldhaar","year":"2011","unstructured":"Feldhaar, H. Bacterial symbionts as mediators of ecologically important traits of insect hosts. Ecological Entomology \n                           36, 533\u2013543 (2011).","journal-title":"Ecological Entomology"},{"key":"11781_CR16","doi-asserted-by":"publisher","first-page":"1473","DOI":"10.1111\/mec.12421","volume":"23","author":"AK Hansen","year":"2014","unstructured":"Hansen, A. K. & Moran, N. A. The impact of microbial symbionts on host plant utilization by herbivorous insects. Molecular Ecology \n                           23, 1473\u20131496 (2014).","journal-title":"Molecular Ecology"},{"key":"11781_CR17","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s00442-015-3327-1","volume":"179","author":"TJ Hammer","year":"2015","unstructured":"Hammer, T. J. & Bowers, M. D. Gut microbes may facilitate insect herbivory of chemically defended plants. Oecologia \n                           179, 1\u201314 (2015).","journal-title":"Oecologia"},{"key":"11781_CR18","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1146\/annurev.ento.43.1.17","volume":"43","author":"AE Douglas","year":"1998","unstructured":"Douglas, A. E. Nutritional interactions in insect-microbial symbioses: aphids and their symbiotic bacteria buchnera. Annual Review of Entomology \n                           43, 17\u201337 (1998).","journal-title":"Annual Review of Entomology"},{"key":"11781_CR19","doi-asserted-by":"publisher","first-page":"14543","DOI":"10.1073\/pnas.2135345100","volume":"100","author":"NA Moran","year":"2003","unstructured":"Moran, N. A., Plague, G. R., Sandstr\u00f6m, J. P. & Wilcox, J. L. A genomic perspective on nutrient provisioning by bacterial symbionts of insects. Proceedings of the National Academy of Sciences \n                           100, 14543\u201314548 (2003).","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"11781_CR20","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1146\/annurev.genet.41.110306.130119","volume":"42","author":"NA Moran","year":"2008","unstructured":"Moran, N. A., McCutcheon, J. P. & Nakabachi, A. Genomics and evolution of heritable bacterial symbionts. Annual Review of Genetics \n                           42, 165\u2013190 (2008).","journal-title":"Annual Review of Genetics"},{"key":"11781_CR21","doi-asserted-by":"publisher","first-page":"21236","DOI":"10.1073\/pnas.0907926106","volume":"106","author":"JA Russell","year":"2009","unstructured":"Russell, J. A. et al. Bacterial gut symbionts are tightly linked with the evolution of herbivory in ants. Proceedings of the National Academy of Sciences \n                           106, 21236\u201321241 (2009).","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"11781_CR22","doi-asserted-by":"publisher","first-page":"375","DOI":"10.1111\/j.1558-5646.2011.01436.x","volume":"66","author":"J Ferrari","year":"2012","unstructured":"Ferrari, J., West, J. A., Via, S. & Godfray, H. C. J. Population genetic structure and secondary symbionts in host-associated populations of the pea aphid complex. Evolution \n                           66, 375\u2013390 (2012).","journal-title":"Evolution"},{"key":"11781_CR23","doi-asserted-by":"publisher","first-page":"323","DOI":"10.1007\/s00442-007-0730-2","volume":"153","author":"J Ferrari","year":"2007","unstructured":"Ferrari, J., Scarborough, C. L. & Godfray, H. C. J. Genetic variation in the effect of a facultative symbiont on host-plant use by pea aphids. Oecologia \n                           153, 323\u2013329 (2007).","journal-title":"Oecologia"},{"key":"11781_CR24","doi-asserted-by":"publisher","first-page":"760","DOI":"10.1098\/rspb.2010.1654","volume":"278","author":"A McLean","year":"2011","unstructured":"McLean, A., Van Asch, M., Ferrari, J. & Godfray, H. Effects of bacterial secondary symbionts on host plant use in pea aphids. Proceedings of the Royal Society of London B: Biological Sciences \n                           278, 760\u2013766 (2011).","journal-title":"Proceedings of the Royal Society of London B: Biological Sciences"},{"key":"11781_CR25","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s00442-016-3603-8","volume":"181","author":"L Ruokolainen","year":"2016","unstructured":"Ruokolainen, L., Ikonen, S., Makkonen, H. & Hanski, I. Larval growth rate is associated with the composition of the gut microbiota in the Glanville fritillary butterfly. Oecologia \n                           181, 1\u20139 (2016).","journal-title":"Oecologia"},{"key":"11781_CR26","doi-asserted-by":"publisher","first-page":"702","DOI":"10.1046\/j.1365-2311.2002.00458.x","volume":"27","author":"CC Nice","year":"2002","unstructured":"Nice, C. C., Fordyce, J. A., Shapiro, A. M. & Ffrench-Constant, R. Lack of evidence for reproductive isolation among ecologically specialised lycaenid butterflies. Ecological Entomology \n                           27, 702\u2013712 (2002).","journal-title":"Ecological Entomology"},{"key":"11781_CR27","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1155\/2012\/242154","volume":"2012","author":"CF Scholl","year":"2012","unstructured":"Scholl, C. F., Nice, C. C., Fordyce, J. A., Gompert, Z. & Forister, M. L. Larval Performance in the Context of Ecological Diversification and Speciation in Lycaeides Butterflies. 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Scanning electron microscopic views of the shell show that the patterning resides in the outer chorion, while the inner vitelline membrane is relatively flat and featureless. We here describe the morphology of the egg and introduce a physical technique, use of a Dynamitron accelerator, to identify and localize elements in the eggshell. Most elements present are represented in the chorion, but sulfur appears restricted to the vitelline membrane. The micropyle is particularly rich in calcium and, in unhatched eggs, phosphorus as well. J. Morphol. 251:140\u2013148, 2002. \u00a9 2002 Wiley\u2010Liss, Inc.<\/jats:p>","DOI":"10.1002\/jmor.1079","type":"journal-article","created":{"date-parts":[[2002,8,25]],"date-time":"2002-08-25T16:25:38Z","timestamp":1030292738000},"page":"140-148","source":"Crossref","is-referenced-by-count":12,"title":["Egg of the Karner Blue butterfly (<i>Lycaeides melissa samuelis<\/i>): Morphology and elemental analysis"],"prefix":"10.1002","volume":"251","author":[{"given":"Elizabeth P.","family":"Nickles","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Helen","family":"Ghiradella","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hassaram","family":"Bakhru","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Arthur","family":"Haberl","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"311","published-online":{"date-parts":[[2001,11,16]]},"reference":[{"key":"e_1_2_1_2_1","doi-asserted-by":"publisher","DOI":"10.1016\/0012-1606(79)90112-X"},{"key":"e_1_2_1_3_1","doi-asserted-by":"publisher","DOI":"10.1016\/0168-583X(89)91078-1"},{"key":"e_1_2_1_4_1","first-page":"1","article-title":"Chorionic sculpturing in eggs of Lycaenidae. 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