Lugardonia paradoxa gen. et sp. nov., a new strobilus from the Anisian flora of Kühwiesenkopf, the Dolomites, Italy and its affinities with emphasis on spore ultrastructure Evelyn Kustatscher a, ⁎, Alan Hemsley b , Johanna H.A. Van Konijnenburg-van Cittert c a Naturmuseum Südtirol, Bindergasse 1, 39100 Bozen/Bolzano, Italy b School of Earth, Ocean and Planetary Sciences, Cardiff University, Cardiff CF10 3YE, UK c Laboratory of Palaeobotany and Palynology, Budapestlaan 4, 3584 CD Utrecht and National Natural History Museum ‘Naturalis’, PO Box 9517, 2300 RA Leiden, The Netherlands abstract article info Article history: Received 26 November 2007 Received in revised form 9 September 2008 Accepted 22 September 2008 Available online 11 October 2008 Keywords: new reproductive organ Pteridophyta in situ spores ultrastructure Middle Triassic Dolomites Italy Lugardonia paradoxa gen. et sp. nov., a new strobilus with in situ spores from the Anisian flora of Kühwiesenkopf, the Dolomites, Italy, is described. These strobili show the general gross morphology of a gymnosperm (e.g. a seed fern) inflorescence with male organs consisting of pollen sacs on short stalks. However, they yield large trilete spores (c. 100 μm in diameter), almost circular in equatorial outline, with a smooth inner layer and a granulate–verrucate outer layer (similar to exospore and perispore in fern spores). These spores suggest a pteridophyte rather than a gymnosperm affinity. Because of either macromorpho- logical characters or spore morphology (based on light microscope (LM), scanning electron microscope (SEM) and transmission electron microscope (TEM) analysis, most plant groups are ruled out as possible parent plants for these strobili. Only the ferns and seed ferns might be considered as potential parent plants. The possible affinities of these strobili within the ferns and seedferns are discussed in detail, with special emphasis on the ultrastructural features of the in situ spores. However, no definite conclusions can be drawn, although a fern affinity appears to be the more likely one based on spore morphology. © 2008 Elsevier B.V. All rights reserved. 1. Introduction The rich Anisian plant deposit located on the western slope of Kühwiesenkopf/Monte Prà della Vacca (Prags/Braies Dolomites) is the subject of ongoing research. The flora is rather rich in specimens and contains a number of land plant taxa belonging to the Lycophyta, Sphenophyta, Pteridophyta, Pteridospermae, Cycadophyta and Coni- ferophyta, some of which have been the subject of a thorough systematic description (Broglio Loriga et al., 2002; Kustatscher, 2004; Van Konijnenburg-van Cittert et al., 2006; Kustatscher et al., 2007), while others are still under study. In addition to taxa with a clear botanical affinity, some problematic fossils occur in the assemblage. Several spore bearing organs consisting of a long (over 200 mm) axis with microsporangia attached in small groups (3–4) have been recovered. These strobili show the general gross morphology of a gymnosperm (e.g. a seed fern) with male organs consisting of pollen sacs on short stalks. However, they yield large trilete spores, thus suggesting a pteridophyte rather than a gymnosperm affinity. Because of the enigmatic combination of gymnosperm macromor- phology and fern-like spores, these strobili are the subject of this study. They will first be described in detail including spore data from LM, SEM and TEM work; then the possible affinities of these strobili will be discussed and comparisons made with some strobili characterised by comparable macromorphology and/or spore ultrastructure. 2. Material and methods The fossils are contained in a basinal succession which has been attributed to the Dont Formation (Senowbari-Daryan et al., 1993; Broglio Loriga et al., 2002). The Anisian outcrops of the Prags/Braies Dolomites have been famous in the paleontological literature since the 19th century, mainly on account of the Anisian (Pelsonian) brachio- pods and ammonoids of the “alpiner Muschelkalk” (Loretz, 1875; Mojsisovics, 1879, 1882; Bittner, 1890) and calcareous algae (Pia, 1937), while only rarely plant remains have been mentioned (e.g. Pia, 1937; Bechstädt and Brandner, 1970). Broglio Loriga et al. (2002) and Kustatscher et al. (2006) give more details on the locality and its age. The specimens have been studied with the aid of a dissecting microscope and, where possible, in situ spore preparations were made. For this purpose, single microsporangia or clusters of micro- sporangia were macerated in Schulze's reagent (KClO 3 and 30% HNO 3 ) and neutralized with 5% NH 4 OH. The sporangia were then separated Review of Palaeobotany and Palynology 156 (2009) 90–97 ⁎ Corresponding author. Fax: +39 0471 412979. E-mail addresses: Evelyn.Kustatscher@naturmuseum.it (E. Kustatscher), hemsleyar@Cardiff.ac.uk (A. Hemsley), Konijnenburg@naturalis.nnm.nl (J.H.A. Van Konijnenburg-van Cittert). 0034-6667/$ – see front matter © 2008 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2008.09.003 Contents lists available at ScienceDirect Review of Palaeobotany and Palynology journal homepage: www.elsevier.com/locate/revpalbo