From early days of Christianity in the British Isles and Ireland, links have existed between churches of western British regions, especially Wales and Cornwall, and Brittany. Migrations from Ireland and western Britain to Brittany brought manuscripts to communities of monks and nuns there, where their styles influenced Breton book decoration, and manuscripts travelled the other direction, from Brittany to Wales and Cornwall. Even after Charlemagne's programme of book production and monastic standardisation, Breton manuscripts retained their decorative style. In a manuscript such as this gospelbook, animal ornament, interlace and other designs resembling British and Irish decoration, coexist with a script developed at Tours and earlier Continental decoration. The symbols of the evangelists are the four winged creatures which appear in the vision of Ezekiel and in the Book of Revelation (man, lion, ox, eagle) and were interpreted by early Christian writers as signs of the gospels. According to Jerome, John's sign was the eagle, as shown here. Different kinds of depictions of them developed early on, including that seen here, combining the animal head with a human body. By the 9th century, such hybrid depictions of the symbols were rarely depicted. In a region such as Brittany, however, which was remote from major centres and often resistant to Carolingian standardisation, the variant, often archaic traditions survived more easily. The serpents in the lower panels of the frame are splendid examples of Breton-style animal interlace.