Miocene Cupressinoxylon from Gokceada (Imbros), Turkey with Protophytobia cambium mining and the study of ecological signals of wood anatomy

Abstract

Premise: The recognition of the Miocene Climate Optimum (MCO) in terrestrial palaeoenvironments of the Eastern Mediterranean is restricted to Lesbos and Lemnos Islands, Greece. This area is significant for its wood microfossils. A recently -discovered fossil wood assemblage from Gokceada (Imbros) Island, Turkey, including tree species similar to the Greek findings, is thought to have an early Miocene age. Here, we revise the age of the latter plant fossiliferous locality, re-evaluate the area for the study of MCO for the terrestrial palaeoecosystems of the Eastern Mediterranean and the nomenclature errors referring to the occurrence of fossil wood. We present the plant-insect-environment interactions using detailed anatomical descriptions, of an extinct conifer and its extinct cambium miner feeding traces observed in its secondary xylem.Methods: Three thin sections were prepared with standard palaeoxylotomical techniques from a small section of the silicified wood; the sections were observed under a light microscope. The anatomy of the conifer and its damage patterns were compared with those of extant and fossil Cupressaceae and Agromyzidae, respectively.Pivotal results: The common anatomical features of the studied wood specimen and Hesperocyparis macrocarpa (Hartw.) Bartel and a shared characteristic (the number of the cross -field pits - a feature we consider of diagnostic value) with Xanthocyparis vietnamensis Farjon & T.H. Nguyen led to its assignment to the Hesperocyparis-Xanthocyparis-Callitropsis clade. The detailed study of the wound scars and anatomical abnormalities, the anatomical-environmental associations, and structural-functional reactions follow the identification of the wood's anatomy sensu Carlquist providing decisive results.Conclusions: Based on the distinctive characteristics presented, we identify our macrofossil as Cupressinoxylon matromnense Grambast, a stem or an extinct lineage of the Hesperocyparis-Xanthocyparis vietnamensis-Callitropsis nootkatensis clade with feeding traces of the fossil cambium miner of the genus Protophytobia Suss (Diptera: Agromyzidae), and anatomical damage and reaction tissue on adventitious shoots. The use of Protopinaceae and Pinoxylon F. H. Knowlton from the eastern Mediterranean are re-evaluated and corrections are provided. The age of the studied plant fossiliferous locality in Gokceada is revised as middle Miocene, allowing the proposal of an eastern Mediterranean MCO hotspot, including Lesbos, Lemnos, and Gokceada (Imbros) Islands.