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In this journal's terms <http://www.biologists.com/copyright_permissions.html#open> they say that articles under the Open option are CC-BY and will be made available immediately ON THEIR WEBSITE. This article, on the website, says "Free" next to the PDF and full text options; other articles, such as doi: 10.1242/dev.101915 do not, and lead me to a pay wall. Therefore, I conclude this item is CC-BY.