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This paper provides a framework for the European history of biological exchange and biological invasion. It distinguishes four main eras of heightened biological exchange. The first came with the introduction to Europe of the neolithic complex of domesticated plants and animals. The second was associated with the increased trade and travel of the pax romana. The third came with increased connections to Asia and the Indian Ocean world in the era of the Crusades. The fourth, well-known to historians, followed from the oceanic navigations of European mariners beginning in the fifteenth century. In all but the last of these, Europe imported far more than it exported in terms of useful species, and most of those exported were earlier imports, mainly from southwest or south Asia.
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