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(Athrotaxoideae, Cupressaceae) from the Upper Cretaceous Raritan Formation, New Jersey, USA

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(Athrotaxoideae, Cupressaceae) from the Upper Cretaceous Raritan Formation, New Jersey, USA Botany A new species of Athrotaxites (Athrotaxoideae, Cupressaceae) from the Upper Cretaceous Raritan Formation, New Jersey, USA. Journal: Botany Manuscript ID cjb-2016-0061.R1 Manuscript Type: Article Date Submitted by the Author: 19-May-2016 Complete List of Authors: Escapa, Ignacio; CONICET-MEF, Paleobotany Gandolfo, Maria;Draft Cornell University, Department of Plant Biology Crepet, William; Cornell University, Department of Plant Biology Nixon, Kevin; Cornell University, L.H. Bailey Hortorium, Plant Biology Section, School of Integrative Plant Science, Cornell University Keyword: Cupressaceae, Athrotaxites, Cretaceous, Raritan Formation, New Jersey https://mc06.manuscriptcentral.com/botany-pubs Page 1 of 39 Botany A new species of Athrotaxites (Athrotaxoideae, Cupressaceae) from the Upper Cretaceous Raritan Formation, New Jersey, USA. Ignacio H. Escapa, Maria A. Gandolfo, William L. Crepet, and Kevin C. Nixon I. H. Escapa. CONICET, Museo Paleontológico Egidio Feruglio, Avenida Fontana 140, 9100 Trelew, Chubut, Argentina. M. A Gandolfo, W. L. Crepet and K. C. Nixon. L.H. Bailey Hortorium, Plant Biology Section, School of Integrative Plant Science, Cornell University, Ithaca, New York Corresponding author: Ignacio Escapa (e-mail: [email protected]) Draft https://mc06.manuscriptcentral.com/botany-pubs Botany Page 2 of 39 Abstract. A new species of anatomically preserved Cupressaceae is described from the Upper Cretaceous Raritan Formation (New Jersey, USA). The fossils are charcolified isolated ovuliferous complexes that were studied by means of a combination of MEB images and Micro-CT, allowing the observation of morphological and anatomical characters. Each ovuliferous complex bears 3-4 anatropous winged seeds, disposed in one row on a thin medial part of the adaxial side of the ovuliferous complex. Based on the combination of characters such as ovuliferous complex morphology, arrangement of vascular tissues and resin canals, seed number and their morphology, orientation and disposition, these fossils are placed within a new species of the fossil genus Athrotaxites . The developmental stage of the specimens is analyzed base on comparisons with living representatives of the subfamily Athrotaxoideae (i.e., Athrotaxis spp.), which supports a post- pollination stage for these fossils. In addition, the new species is compared with other extant and extinct representatives of basal cupressaceous subfamilies. This new record from the Upper Cretaceous sediments of New Jersey furtherDraft supports a wider distribution of the subfamily Athrotaxoideae during the middle part of the Mesozoic, as it has been also noted for other basal representatives of the family Cupressaceae. Key words: Cupressaceae. Athrotaxoideae. Athrotaxites . Cretaceous. Raritan Formation. New Jersey https://mc06.manuscriptcentral.com/botany-pubs Page 3 of 39 Botany Introduction The well-preserved, charcoalified angiosperm remains from the Turonian Raritan Formation are comprehensively known through numerous studies during the last two decades (e.g., Crepet et al. 1992, 2005, 2013; Herendeen et al. 1993, 1994; Nixon and Crepet 1993; Crepet and Nixon 1994; Gandolfo et al. 1998 a, 1998 b, 2002, 2004; Hermsen et al. 2003; Martínez-Millán et al. 2009). Fossil diversity at the Old Crossman Clay Pit locality (Sayreville, New Jersey) also includes ferns (Gandolfo et al. 1997, 2000) and gymnosperms (e.g. Miller 1985), that, in contrast to angiosperm fossils, have been scarcely studied. Three-dimensionally preserved gymnosperms from this formation have been known since the beginning of the 20 th century; however, only two species of pinaceous leaves ( Prepinus crossmanensis and P. quinquefolia ) and the pollen cone Amboystrobus cretacicum (Gandolfo et al. 2001) have been described in detail on the basis of charcoalified specimens. The conifer family Cupressaceae sensu lato (including the paraphyletic former Taxodiaceae) has been proposed based on numerous morphologicalDraft and molecular phylogenetic analyses (Eckenwalder 1976; Hart 1987; Price and Lowenstein 1989; Chase et al . 1993; Brunsfeld et al. 1994; Stefanovic et al. 1998; Gadek et al. 2000; Kusumi et al. 2000; Farjon 2005). Twenty genera and 142 extant species are recognized in Cupressaceae, which is the most diverse conifer family in terms of genera. Seven extant subfamilies are currently recognized (Gadek et al. 2000). Interestingly, there is a strong distinction between the Northern and Southern Hemisphere lineages of Cupressaceae sensu lato (see Leslie et al. 2012) . Nevertheless, this distinction is less definite if fossil and extant species are simultaneously considered. For instance, the contrast between the restricted distribution of extant Cunninghamia (Liu 1966) and the widespread occurrence and notable diversity of the subfamily Cunninghamioideae during the Mesozoic and Cenozoic has been recognized (Stockey et al. 2005; Escapa et al. 2008; Serbet et al. 2013; Shi et al. 2014). The extremely low extant diversity and a similar distribution pattern is also shared with the subfamilies Taiwanioideae and Athrotaxoideae, which occur in a pectinate arrangement together with Cunninghmamia at the base of the Cupressaceae phylogenetic tree (Leslie et al. 2012) . Extant representatives of Cupressaceae subfamily Athrotaxoideae comprise three species: Athrotaxis selaginoides, A. laxifolia, and A. cupressoides ; all of which are distributed in temperate https://mc06.manuscriptcentral.com/botany-pubs Botany Page 4 of 39 rainforests and alpine vegetation in Tasmania (Cullen and Kirkpatrick 1988; Farjon 2005). However, A. laxifolia is now considered to be an interspecific hybrid between the other two species, based on morphological (Isoda et al. 2000) and genetic evidence (Worth et al. 2016). Molecular models suggested a Neogene-Paleogene age of divergence for the Ahtrotaxis crown group, while the stem age has been estimated as Jurassic-Late Cretaceous (Leslie et al. 2012; Mao et al. 2012; Yang et al. 2012). The fossil record of Athrotaxoideae has been extended to the Early Cretaceous, either with fossils assigned to Athrotaxis or to the fossil genus Athrotaxites (Archangelsky 1963; Miller and LaPasha 1983; Stockey et al. 2005). Fossil evidence for this group includes a small number of Cretaceous records in both hemispheres, together with a more diverse Cenozoic record (see Hill et al. 1993; Hill and Brodribb 1999). In this contribution, we report the occurrence of athrotaxoid fossils from the Upper Cretaceous Raritan Formation (New Jersey, USA) based on the presence of charcoalified ovuliferous complexes that belong to the subfamily AthrotaxoideDraftae . The combined use of Scanning Electron Microscopy (SEM) and Micro-CT Scan allowed the observation of detailed anatomical and morphological features supporting the taxonomic placement of these fossils within the fossil genus Athrotaxites and the erection of a new species, probably representing a stem component within the subfamily Athrotaxoideae. Materials and methods Geological setting and paleontological background The specimens described in this study were collected from the Old Crossman Clay Pit locality, Sayreville (New Jersey, USA), in outcrops of the South Amboy Fire Clay, a member of the Raritan Formation. The Raritan is part of fluvial sediments composing the Atlantic Coastal plains (see Petters 1976). Based on regional lithostratigraphy, the sediments of the Raritan Fm. exposed in New Jersey are divided into five members: (i) the Raritan Fire Clay, (ii) Farrington Sand, (iii) Woodbridge Clay, (iv) Sayreville Sand, and (v) South Amboy Fire Clay. The age of this member is calculated as Turonian (Upper Cretaceous, 90–94 MY, see Harland et al. 1989) based on stratigraphic and palynological data. Since end of the 19 th century, the Raritan Fm. is considered of special interest https://mc06.manuscriptcentral.com/botany-pubs Page 5 of 39 Botany because of its paleobotanical and palynological content (e.g. Newberry 1895). Subsequent studies led to the first compendium of the flora by Berry (1911) and later revisions (e.g., Dorf 1952). Notably, during the last 25 years, the majority of the paleobotanical studies on its paleoflora were focused on its charcoalified specimens which are highly diverse. The flora is dominated by angiosperms, while several conifers seem to be well-represented but not as abundant; the pteridophytes also are represented by a few taxa. The angiosperms are represented by flowers, fruits, and seeds of various phylogenetically disparate groups of magnoliid, monocots and eudicots (e.g., Crepet et al. 1992, 2005; Herendeen et al. 1993, 1994; Nixon and Crepet 1993; Crepet and Nixon 1994, 1998; Gandolfo et al. 1998 a, 1998b, 2001, 2002, 2004; Crepet 2000; Zhou et al. 2001; Hermsen et al. 2003; Martínez- Millán et al. 2009; Crepet et al. 2013). Study Technique Fossil specimens are charcoalified, with three-dimensional morphology and excellent anatomical details. External features wereDraft studied using scanning electron microscopy (SEM). Specimens were mounted on stubs and sputter-coated with gold/palladium in preparation for examination with a Hitachi 4500 SEM. In addition, the internal anatomy of one specimen was imaged using a Micro-CT (Xradia model xrm-500) at Cornell University (http://www.ct.cornell.edu/cct/Cornell_CT.html), which provided serial sections of the fossil that are the bases for the three-dimensional models (OsiriX 64 bit DICOM viewer). The use of this technique for the anatomical study of fossils has been increasingly used
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