The beetles Rhopalomesites Tardyi, Eurynebria complanata, and Otiorrhynchus auropunctatus also belong to this fauna, as also the Earthworms Allolobophora veneta and A. Georgii, and the Millipede Polydesmus gallicus.
It will be evident to every one from these few instances of Lusitanian species, that somewhere in South-western Europe and North-western Africa, and also, perhaps, in a larger now submerged western land-area, there existed an active centre of development, from which animals spread in all directions.
If the presence of Platyarthrus in North-west Africa proves that the Straits of Gibraltar had come into existence after its southward migration, it also suggests that the ancestral home of this woodlouse was in the Spanish peninsula. Whether this supposition is correct or not, does not affect the Straits of Gibraltar problem, for in a migration northward into Spain from Morocco a land-connection would be equally necessary. Almost every group of vertebrates and invertebrates furnishes instances of species which must have crossed the Straits on dry land. Many naturalists have come to this conclusion, and have clearly expressed their views on the subject. At the commencement of the present period, says Mr. Bourguignat (p. 354), the north of Africa was a peninsula of Spain, the Straits of Gibraltar did not exist, and the Mediterranean communicated by the Sahara with the Atlantic.
The faunas of North-west Africa and the south-western portion of our continent are so closely related, that an uninterrupted intercourse by land must have existed for a very long period. The Mediterranean, however, throughout the Tertiary period—at any rate since miocene times—must have had almost constant communication with the Atlantic. According to Professor Suess, this was the case. The Atlantic was joined with the Mediterranean across the valley of the Guadalquivir during the Miocene Epoch, so that Andalusia must have belonged to North Africa in those days. The Straits of Gibraltar are supposed to have been formed in the next epoch. I have already expressed my disagreement with that theory from a zoogeographical point of view. The old Guadalquivir connection probably persisted much longer,—though interrupted by temporary periods of a partial retreat—so as to uncover sufficient land to allow of an interchange during miocene as well as pliocene times between the European and North African faunas. It is in this way, perhaps, that some of the members of the Alpine fauna have reached Spain by way of Corsica, Sardinia, and North-western Africa, and vice versâ. The Balearic Islands were then connected with Spain; and we find there many curious survivals which have long ago become extinct on the mainland.
That the Straits of Gibraltar are only of recent formation has been suggested on zoogeographical evidence by Bourguignat, Simroth, Kobelt, and many others. Dr. Kobelt believes that the former land-connection between the south of Spain and Morocco was much wider than is generally assumed, and that the coast-line stretched from Oran in Algeria straight across to Cartagena in Spain (b, ii., p. 228).
My allusions to the lands lying beyond the Lusitanian province, refer chiefly to the Canary Islands and Madeira. Whatever doubts Dr. Wallace had on the subject of their former connection with Morocco, it cannot be denied that they used to be of much larger extent, especially in miocene and pliocene times. It seems extremely probable that these islands formed part of the mainland of North Africa until comparatively recently, and that they are the last traces of a sunken continent which united Africa and South America. A discussion of this problem, however, must be deferred, as it is a complicated one, and one which would lead me altogether outside the scope of this little volume. I hope I shall have an opportunity to publish my views on this subject before long, meanwhile the reader must content himself with this mere statement.
During the greater portion of the Miocene, and I think for part of the Pliocene Epoch too, the advance of the Lusitanian species eastward was barred on the continent of Europe by an arm of the sea which stretched northward along the Rhone valley from the Mediterranean. The Lusitanian forms which originated in Southern Spain were able to travel east during these times by way of North-west Africa, Sicily, Southern Italy, and Greece; it is possible that some may have reached the Alps in this manner, and Eastern Europe generally. That the Lusitanian centre was never a very active one compared with, for instance, the Oriental is indicated by many distributional facts. It is difficult to understand, however, why the Oriental species, on the whole, have migrated so far west, while few Lusitanians have gone very far east. This seems to have been noted particularly in the case of the flora. Mr. Bonnet drew attention to the fact that in Tunis there are none of the absolutely characteristic plants of Morocco and Spain, while the Oriental flora is represented by a good many species. Lusitanian species have spread chiefly southward into North Africa, and northward into France, the British Islands, and even Scandinavia. As I have mentioned in the third chapter, there are a good many species of Lusitanian origin in the British Islands. However, we have only a mere remnant of what we ought to have, had the climate been less trying. It is probable, too, that the submergence destroyed a good many plants and the insects dependent on them. That the Lusitanian fauna is very ancient in the British Islands is proved by the fact of the discontinuous distribution of so many species. A greater number survived in Ireland than in England.
Fig. 20.—The Strawberry-tree (Arbutus unedo) in its native habitat in the south-west of Ireland. (From a photograph by Robert Welch.)