Ilex geissertii sp. n. (Aquifoliaceae), a fossil ancestor of Ilex sect. Ilex in the upper Miocene and Pliocene of Europe

Leaf remains of Ilex L. sect. Ilex known from the Pliocene of Auenheim (Alsace, France) and Frankfurt a. M. - Niederrad (Hesse, Germany) as Ilex aff. cornuta Lindley fossilis Geissert and Ilex aquifolium L. fossilis Engelhardt, respectively, are re-interpreted as representing a new species Ilex geissertii Kvaček, Teodoridis et Wang Qing, sp. nov. Similar leaf fossils occur also in the upper Miocene of the Netherlands (Brunssum), France (Murat), and the Pliocene of Italy (Valdarno Superiore). Although these fossil records were considered either direct ancestors of I. aquifolium L. (Near-East, Europe, N Africa) or allied to I. cornuta Lindley and Paxton (China and Korea), both the mentioned extant species differ in the leaf morphology. Close affinities of I.geissertii to several extant representatives of Ilex subsect. Ilex (= Oxyodontae Loesener nom. illegit.) are indisputable and stress a common origin of this group of the hollies known already from the Oligocene in Europe (Ilex castellii Kvaček et Walther). Several extinct elements of the European late Miocene and Pliocene flora accompanying this new holly (e.g., Ginkgo, Taxodium, Fagus kraeuselii, Parrotia, Craigia, and Trichosanthes) share similar former phyllogeography in having their modern nearest relatives outside Europe.