In 2013 I had an academic paper accepted by the Journal of Late Antique Religion and Culture (JLARC). The journal is open access and you can read it here (link to pdf in the right hand column). I found that the manuscript animals were copied from manuscript to manuscript by closeted scribes, and were not based on real animals (e.g. lynxes, cats, wolves).
Species: The manuscript images were all derivative and made up a coherent, although unrealistic tradition of depicting lions (Panthera leo) not lynxes (Lynx lynx), wolves (Canis lupus) or cats (Felis catus; Felis sylvestris).
Source: Some of the oldest manuscripts in Britain: The illuminated gospels.
Date: 650-1000 A.D.
Highlights: One lion, that from the Lichfield (St Chad) Gospel (above) provoked a great deal of interest. It only had the stub of a tail and the scribe’s style made its body looked speckled. However in every other respect the lion was drawn in a derivative way to the mainstream tradition, and the answer is probably that the scribe simply forgot to paint the rest of the lion’s tail.
The majority of the very earliest surviving manuscripts in Europe are religious in nature, and the most important of these manuscripts for medieval audiences are arguably the illuminated gospels. These manuscripts contain the first four books of the New Testament of the Bible (the “synoptic gospels”): Matthew, Mark, Luke and John.
But the illuminated gospel manuscripts are far more than just texts. Each manuscript is a work of art: The handwriting is painstaking. Capital letters starting paragraphs are huge and artistically decorated with patterns or flowers or animals.
Each manuscript usually also contained at least four full-page illustrations. These are called the “evangelist portraits” and are pictures of the evangelists said to have written each gospel. Sometimes the evangelists are also accompanied by their “evangelist symbol”. St John is accompanied by an eagle, St Matthew is accompanied by a man or an angel, St Luke is accompanied by a calf and St Mark is accompanied by a lion.
My aim with these manuscripts was to look at all the early “insular” manuscripts from the island of Britain and look at the depictions of lions. What could have inspired the authors drawings? There were very few lions between the last ones shipped to Britain to battle in Roman Amphitheatres in the fifth century and the opening of the menagerie at Woodstock Palace in the twelfth century.
Some of the findings were quite surprising. British evangelist portraits quite often contain just the evangelist’s animal, and do not show them at all.
The British lions mainly exhibited certain characteristics. They all faced in the same direction, were painted in similar colours and had their tongues stuck out. Contrary to continental examples, most of the British symbols lacked wings. The reason this is important is simple: the fact that all the lions share these characteristics suggest they were copied from one manuscript to another. There is nothing to suggest they were inspired by real-life animals.
You can read the study in full here.