Neutron tomography of Austrosequoia novae-zeelandiae comb. nov. (Late Cretaceous, Chatham Islands, New Zealand): implications for Sequoioideae phylogeny and biogeography

Abstract

The Tupuangi Flora of the Chatham Islands, New Zealand, reveals a south polar forest ecosystem, and important biogeographical links between eastern and western Gondwana. We employed neutron tomography (NT) to image fossil Cupressaceae seed cones from the Upper Cretaceous (Cenomanian) strata of the Tupuangi Formation. This technique facilitated the non-destructive ‘virtual extraction’ of three-dimensional, coalified specimens, whilst they were still embedded within a large volume of supporting silicate sedimentary rock. This study is the first reported application of NT in palaeobotanical taxonomy, and the combination of virtual and manual extraction techniques enabled a more complete treatment than would otherwise be possible if taxonomic data were limited to only one of these approaches. The seed cones were identified as Austrosequoia novae-zeelandiae (Ettingshausen) Mays & Cantrill comb. nov. In this case, NT data supplemented the compression fossil data by providing details such as the three-dimensional measurements of the gross morphology, and accurate estimations of bract-scale complex number. Furthermore, this technique appears to show promise in differentiating between organic compounds within an individual specimen. However, anatomical details and fine-scale morphology were indiscernible due to present limitations in spatial resolution. Austrosequoia novae-zeelandiae is interpreted as a stem group of Sequoioideae; it shares synapomorphic seed cone characters with extant sequoioids (e.g. Sequoia and Sequoiadendron), and plesiomorphic stomatal structures and leaf morphology. Abundant epiphyllous fungi (Plochmopeltinites sp.; Microthyriaceae) were also identified on the leaf cuticles of A. novae-zeelandiae. The high abundance of Austrosequoia in the Tupuangi Flora supports a cupressaceous floral province at south polar latitudes during the early Late Cretaceous. Furthermore, this stem group of Sequoioideae in eastern Gondwana during the early Late Cretaceous suggests an alternative, south-to-north dispersal route of sequoioids before the final continental separation of eastern and western Gondwana.