CORE Metadata, citation and similar papers at core.ac.uk Provided by Repositorio Universidad de Zaragoza Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 1–12, 2018 Agora Paleobotanica The extinct tree fern Tempskya Corda from the Albian of Spain: palaeophytogeographical and palaeoenvironmental implications Luis M. Sender1,2*, Harufumi Nishida2,3 and Jose´ B. Diez4 1 A´ rea de Paleontologı´a, Facultad de Ciencias Geolo´gicas, Universidad de Zaragoza, 50002 Zaragoza, Spain. Email: [email protected] 2 Faculty of Science and Engineering, Chuo University, 1-13-27 Kasuga, Bunkyo, Tokyo, Japan. 3 Graduate School of The University of Tokyo, Hongo, Bunkyo, Tokyo, Japan. 4 Departamento de Xeociencias Marin˜as e Ordenacio´n do Territorio, Facultade de Ciencias do Mar, Universidade de Vigo, 36310 Vigo, Spain. *Corresponding author ABSTRACT: New evidence of the extinct tree fern of the genus Tempskya Corda from Albian deposits in western Eurasia (northeastern Spain) is presented. These plant fossil remains consist of several silicified false trunks measuring up to 1.20 m long, some of which still preserve the apex. Rhizomes and petioles are more abundant in the apical zone of the false trunk. Some false trunks preserve charcoalified tissues that can be interpreted as evidence that palaeo-wildfires affected the false trunks several times from mostly the same direction. Sedimentological evidence suggests that the fern habitat was close to coastal, tidally influenced environments. These new fossils from the Albian of Spain fill the chronological and distributional gap of Tempskya that existed in Eurasia during the Early Cretaceous. Temporal and spatial changes in Tempskya distributions are proposed on several palaeogeographical maps. KEY WORDS: charcoal, early cretaceous, Eurasia, palaeoenvironment, palaeophytogeography, wildfires. The genus Tempskya Corda is a group of extinct ferns com- are generally represented in Eurasia by silicified fragments posed of species that are characterised by a false trunk, which of false trunks that have been recorded in the Lower and consists of numerous intermingled rhizomes and a large ‘Middle’ Cretaceous from the Valanginian–Hauterivian number of adventitious roots coming out from the rhizomes. (Barale & Viera 1989) and Hauterivian–Barremian (Puente- This false trunk can reach up to 50 cm in diameter and 6 m Arauzo et al. 2014) in Spain, the ‘Wealden Beds’ in England high (Taylor et al. 2009). Due to the unique morphology of (Seward 1894; Stopes 1915; Austen & Batten 2011), Germany this false trunk, Tempskya represents an extinct monotypic (Jung 1983) and France (Corda 1845; Carpentier 1923), and family Tempskyaceae Read & Brown ex L.C.A. Martı´nez, from the Cenomanian in the Czech Republic (Corda 1845; 2015. Velenovsky 1888). Nevertheless, this taxon has not previously However, the systematic affinity of the family has remained been recorded from Albian deposits in western Eurasia. unclear to date because of the lack of attached leaves and Here, we describe the first Albian remains of Tempskya reproductive organs (Tidwell & Ash 1994). Recently, Martı´nez false trunks in western Eurasia from middle to upper Albian & Olivo (2015) emended the Tempskyaceae family based deposits in Spain. These consist of several false trunks of on exceptionally well-preserved silicified false trunks, which differing sizes, some of which also present possible evidence included possible frond segments and attached sori with of palaeo-wildfire damage. These new findings fill the temporal sporangia containing spores. The characters of these reproduc- gap of this genus in Eurasia, providing new evidence for tive remains allowed the authors to attribute this family to the inferring the palaeoenvironment and distribution patterns of Cyatheales. Some other Tempskya remains have also been this genus during the Cretaceous. compared to the Anemiaceae, although no direct evidence supporting this affinity exists (Tidwell & Ash 1994). The genus Tempskya has been recorded exclusively in 1. Material and methods Cretaceous deposits from the Valanginian (Barale & Viera 1989; Martı´nez & Olivo 2015) to the Santonian (Nishida 1.1. Geographical and geological setting 1986). Records of Tempskya are found in both Lower and The fossil plants studied in this paper were collected at the Upper Cretaceous deposits from the Laurasia domain, includ- ‘Carretera-1’ fossil site near the village of Escucha in Teruel ing North America, but the genus has also been reported from Province (Arago´n, NE Spain; Fig. 1). This village is situated the Lower Cretaceous of Argentina and Australia within the 70 km N of Teruel city and 5 km E of Utrillas village, which Gondwana realm (see Section 2.4). The remains of this fern is a well-known reference locality for both Albian geology #*" #%#!6 2018 The''$&***!%#%#% Royal Society of Edinburgh.")%&'+#&'%""'%##" ' ' &( doi:10.1017/S1755691018000543 ''#'!%#%'%!&#(&) ' ''$&***!%#%#%'%!&''$&##% 2 LUIS M. SENDER ET AL. Figure 1 (A) Geographical location of the plant fossil site. (B) Geological map of the fossil site near Escucha village. Figure 2 (A) Synthetic stratigraphic section of part of the Escucha and Utrillas Formations, indicating deposi- tional environments and sub-environments (modified from Villanueva-Amadoz 2009). The stratigraphic level bearing Tempskya is indicated by an arrow. (B) Fossil site image with the layer containing Tempskya indicated by an asterisk. See a hammer for scale (28 cm long), to the left above the asterisk. #*" #%#!''$&***!%#%#%")%&'+#&'%""'%##" ' ' &(''#'!%#%'%!&#(&) ' ''$&***!%#%#%'%!&''$&##% TEMPSKYA FROM THE ALBIAN OF SPAIN: IMPLICATIONS 3 and palaeobotany in the region. The fossil locality is in the The false trunks measure up to 1.20 m long in the largest Cuencas Mineras district, and the stratigraphic level contain- specimen (Fig. 3a), with an average diameter of 10 cm, and ing Tempskya (Fig. 2a) belongs to the Upper Member of the consist of thick root mantles (adventitious roots) and embedded Escucha Formation (Aguilar et al. 1971). This formation is rhizomes (Figs 3, 4). Most specimens are somewhat compressed mainly a continental detrital unit that consists of alternate laterally, resulting in an oval outline in cross-sectional view of sandstones and clays intercalated with coal seams and sporadic the false trunk (Fig. 4a, b). marine limestones, which were deposited in a tidally influenced The rhizomes are clearly visible on the surface of both delta-estuary system (Pardo 1979; Pardo & Villena 1979; Querol medial and apical parts of the false trunks (Fig. 3b, d), but 1990; Pardo et al. 1991; Querol et al. 1992; Salas et al. 2001). are often protruded or obscured by root mantles in the basal The Escucha Formation in this area is limited at the base part (Fig. 3c). The rhizomes are distributed randomly in the by the carbonated marine deposits of the Villarroya de los false trunks (Fig. 3b–e) and proceed distally from the base Pinares Formation (Aptian in age) and is overlain by sandy to the apex. Interval dichotomisation of rhizomes is ca.5cm fluvial deposits of the Utrillas Formation (middle–late Albian). on average, with the next order of rhizomes also dividing The most recent palynological studies in the area, where the dichotomously at approximate intervals of 1.5–2 cm (Fig. 3d, e). remains of Tempskya were found, indicate a middle to late In cross section, rhizomes are oval-shaped and arranged Albian age for the Upper Member of the Escucha Formation randomly in the false trunks (Fig. 4). The leaf traces depart (Villanueva-Amadoz 2009). The lithology of the stratigraphic from the dorsal side of the rhizomes, either singly or as part interval in the Escucha locality bearing the studied fossils of aggregations of several traces, with shorter internodes in exhibits white cross-bedded sandstones intercalated with shales most rhizomes (Figs 5a, b, d). One specimen (MPZ2017/540) and grey-coloured clays (Fig. 2b) deposited in a fluvio-tidal shows leaf petiole scars arranged in a definite interval of nearly environment (Pardo et al. 1991; Querol et al. 1992). 0.7 cm on the ventral side of the rhizome (Fig. 3f). Histological details are best preserved in specimen MPZ2017/532 (Fig. 5a), 1.2. Studied material although the preservation is still insufficient to observe tissue Fragments of permineralised false trunks (24 pieces) possibly details. represent 14 individual plants. The largest specimen measures Thin sections of the rhizomes show their structure from the 1.20 m long (Fig. 3a). All false trunks are preserved as silicified centre outwards, consisting in this order of the pith, the vascular remains, although they have undergone both recrystallisation cylinder, the cortex and the epidermis. The different tissues can and weathering, causing some alteration of internal structures be recognised as concentric areas preserved differently both in (Fig. 4a). In addition, some samples exhibit compressional colour and in structure (Fig. 5b, d). The cortex is composed deformation. Three false trunks contain some charcoalified of the outer layer, which is preserved as a pale-coloured zone remains preserved within the surrounding silicified tissues that is probably parenchymatous, and the inner layer, which (Figs 4b, c, 6, 7). consists of a dark-coloured band and is possibly sclerenchymatous Fossils were first cleaned with fresh water and then five (Fig. 5b). The vascular cylinder is amphiphloic, with a central fragments of fossil trunks were cut transversally in order to