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To Link and Cite This Article: Doi: 10.5710/AMGH.23.06.2020.3333 Submitted: January 13th, 2020 – Accepted: June 23th, 2020 – Published online: June 28th, 2020 To link and cite this article: doi: 10.5710/AMGH.23.06.2020.3333 1 A NEW SPECIES OF BRACHYPHYLLUM FROM THE CRATO FORMATION 2 (LOWER CRETACEOUS), ARARIPE BASIN, BRAZIL 3 4 MARIA EDENILCE P. BATISTA1,2*, LUTZ KUNZMANN3, ARTUR A.A. SÁ4,5, 5 ANTÔNIO ÁLAMO F. SARAIVA2, MARIA IRACEMA B. LOIOLA6 6 1Programa de Pós-Graduação em Ecologia e Recursos Naturais, Departamento de Biologia, 7 Universidade Federal do Ceará, Av. Mister Hull, CEP 60455-900, Fortaleza, Brazil. 8 2Departamento de Ciências Biológicas, Universidade Regional do Cariri, R. Cel. Antônio 9 Luis, CEP 63105-000, Crato, Brazil. 3Senckenberg Natural History Collections, 10 Königsbrücker Landstr. 159, 01109 Dresden, Germany. 4Departamento de Geologia, 11 Universidade de Trás-os-Montes e Alto Douro, Quinta de Prados, 5000-801 Vila Real, 12 Portugal. 5Centro de Geociências, Universidade de Coimbra-Polo II, 3030-790 Coimbra, 13 Portugal. 6Departamento de Biologia, Universidade Federal do Ceará, Av. Mister Hull, CEP 14 60455-900, Fortaleza, Brazil. 15 20 pag. (text+references); 5 figs. 2 tables 16 BATISTA ET AL.: NEW BRACHYPHYLLUM FROM THE ARARIPE BASIN, BRAZIL 17 permineralized 18 Running Header: We describe a new species of the fossil genus Brachyphyllum from the 19 Crato Formation, Araripe Basin, Brazil. 20 21 Corresponding author: Maria Edenilce Peixoto Batista, e-mail [email protected] 22 23 24 25 26 27 Abstract. The Araripe Basin, located on the Northeast of Brazil, contains many fossils. 28 Among these, the gymnosperms stand out, especially species of the fossil taxon 29 Brachyphyllum. It encompasses conifer shoots with helically arranged scale-like, adpressed 30 leaves whose length does not exceed the width. This foliage morphotype was widely 31 distributed during the Mesozoic and species accommodated in this genus can be attributed to 32 different families if their anatomical characters are taken into consideration. In the Araripe 33 basin, specifically in the Crato and Romualdo formations, B. obesum is frequently found and 34 was attributed to the family Araucariaceae based on leaf epidermal and xylem characters. In 35 this work, we describe the new species from the Crato Formation (Lower Cretaceous), 36 Brachyphyllum sattlerae sp. nov., based on the following morphological and anatomical 37 features: submain shoots with lateral branches oppositely disposed and almost exclusively 38 decussately arranged leaves; scarious leaf margins; rugose abaxial leaf surfaces; non-papillate 39 epidermal cells; stomata distributed in longitudinal rows except close to the leaf margins and 40 scarce at the leaf base; stomatal openings obliquely oriented; and hypodermis with a sinuous 41 pattern around the substomatal chamber. Some of these features are distinctive of 42 Araucariaceae, which indicates that B. sattlerae sp. nov. also belongs to this family. 43 44 45 Key words. Araucariaceae. Cretaceous. Conifer. Epidermal micromorphology. 46 47 Resumen. UNA NUEVA ESPECIE DE BRACHYPHYLLUM DE LA FORMACIÓN CRATO 48 (CRETÁCICO INFERIOR), CUENCA ARARIPE, BRASIL. La cuenca de Araripe, ubicada 49 en el noreste de Brasil, contiene mumerosos registros fósiles. Entre estos, se destacan las 50 gimnospermas, especialmente las especies del género fósil Brachyphyllum. Se registran ramas 51 de coníferas con hojas dispuestas helicoidalmente en forma de escamas, adheridas, cuya 52 longitud no excede el ancho. Este tipo foliar se distribuyó ampliamente durante el Mesozoico 53 y las especies asignadas en este género pueden atribuirse a diferentes familias si se tienen en 54 cuenta sus caracteres anatómicos. En la cuenca de Araripe, específicamente en las 55 Formaciones Crato y Romualdo, B. obesum se encuentra con frecuencia y fue atribuida a la 56 Familia Araucariaceae por sus caracteres epidérmicos y el xilema de las hojas. En este 57 trabajo, describimos una nueva especie recuperada en la Formación Crato (Cretácico Inferior), 58 Brachyphyllum sattlerae sp. nov., basada en las siguientes características morfológicas y 59 anatómicas: brote subprincipal con ramas laterales con disposición opuesta y hojas casi 60 exclusivamente dispuestas de forma decusada; márgenes de las hojas escariosas; las 61 superficies abaxiales de las hojas son rugosas; las células epidérmicas sin papiladas; estomas 62 distribuidos en hileras longitudinales excepto cerca de los márgenes de las hojas siendo 63 escasos en la base de la hoja; aberturas estomáticas orientadas oblicuamente; e hipodermis 64 con un patrón sinuoso alrededor de la cámara subestomática. Algunas de estas características 65 son distintivas de Araucariaceae, lo que indica que B. sattlerae sp. nov. pertenece a esta 66 familia. 67 68 69 Palabras clave. Araucariaceae. Cretácico. Coníferas. Micromorfología epidérmica. 70 71 72 73 74 75 76 77 THE Araripe Basin in northeastern Brasil has been known for a long time for its exquisitely 78 preserved fossils recovered from several formations. The basin has already been included 79 among the top ten of the fossil lagerstätten in the world (Maisey, 1991; Martill, 1990). One of 80 its units, the Lower Cretaceous Crato Formation, is considered one of the best windows to a 81 northern Gondwanan Cretaceous ecosystem (Martill et al., 2007). The Crato fossil beds are a 82 classical konservat lagerstätte containing excellently preserved vertebrates, invertebrates and 83 plants. The Crato taphoflora is rather diverse and includes charophytes, sphenophytes, 84 lycophytes, monilophytes, gnetaleans, conifers, pteridospermophytes, cycadophytes, and 85 angiosperms, (Bernardes-de-Oliveira et al., 2007; Fanton, 2007; Martill et al., 2007; Mohr et 86 al., 2000; Mohr et al., 2007). As low latitude fossil flora, it lacks typical mid to high latitude 87 groups such as gingkoaleans. 88 Conifers have been described from megafossil remains and pollen assemblages. 89 Among megafossils, representatives of the families Araucariaceae, Cheirolepidiaceae, and 90 Podozamitaceae were determined (Duarte, 1985, 1993; Kunzmann et al., 2004, 2006; Mohr et 91 al., 2007, 2012; Sucerquia et al., 2015; Batista et al., 2017), while Cupressecae and Pinaceae 92 were reported from pollen assemblages (Lima, 1979; Batten, 2007; Portela et al., 2014). 93 Species of Araucariaceae and a considerable number of species of Cheirolepidiaceae are 94 characterized as large canopy-forming trees (Bernardes-de-Oliveira et al., 2007; Martill et al., 95 2007; Barral et al., 2016). Foliage shoots of the taxon Brachyphyllum Lindl. et Hutton ex 96 Brongn. emend. T.M.Harris are the most frequent plant fossils of the Crato flora (Duarte, 97 1985; Kunzmann et al., 2004; Batista et al., 2017). However, in contrast to the number of 98 specimens reported, the diversity of fossil species determined from the Crato flora is 99 comparatively low. Only two species B. obesum Heer and B. castilhoi L.Duarte have been 100 described to date (Batista et al., 2017). While the majority of the Brachyphyllum shoots are 101 ascribed to Among B. obesum, the latter species is quite rare (Duarte, 1985, 1993; Martill et 102 al., 2007). 103 Representatives of the fossil genus Brachyphyllum have been reported from all 104 continents during the Mesozoic Era (Traverso, 1968; Ash, 1973; Raab et al., 1986; Yabe & 105 Kubota, 2004; Barale & Appert, 2007; Du et al., 2013; Karakitsios et al., 2015). It is 106 characterized by helically arranged,scale-like adpressed leaves, whose total length (length of 107 the free part and length of the cushion) is the same or less than the width of the leaf cushion 108 (Kendall, 1947; Harris, 1979). According to the concept of Harris (1979), who gave an 109 emended diagnosis, Brachyphyllum is an artificial morphotype. If anatomical characters, such 110 as leaf epidermis and wood structure, are preserved and taken into consideration for 111 taxonomic purposes individual species show clear affinities to different families. To date, 112 Brachyphyllum species are affiliated to Araucariaceae, Cheirolepidiaceae, Podocarpaceae, 113 and Cupressaceae (Hollick & Jeffrey, 1906; Kendalll, 1947; Harris, 1979; van der Ham et al. 114 2003; Kunzmann et al., 2004; Batista et al., 2017). For instance, the aforementioned B. 115 obesum was repeatedly assigned to Araucariaceae (Kunzmann et al., 2004; Batista et al., 116 2017), whereas the systematic relationships of B. castilhoi are still indefinite. This taxonomic 117 question can only be resolved if appropriately preserved fossil material will be available for 118 micro- and ultrastructural studies. 119 In this paper, we analyze the morphology and epidermal structure of peculiar 120 Brachyphyllum specimens from the Crato Formation whose features exclude an attribution to 121 both B. obesum and B. castilhoi. Based on a distinct set of characters we propose the new 122 species of Brachyphyllum sattlerae sp. nov., which most likely belongs to the Araucariaceae. 123 Therefore, our result contributes to better understand the diversity and frequency of conifers 124 in a paleo-equatorial environment of Gondwana. 125 126 GEOLOGICAL SETTING 127 The Araripe Basin is located upon the central part of a continental collision belt called 128 the Proterozoic Borborema Tectonic Province (Assine, 1992, 2007; Valença et al., 2003). 129 Geographically, this Mesozoic basin is situated in the border areas of Ceará, Pernambuco and 130 Piauí states. It is the most extensive intracratonic basin of northeastern Brazil, covering an 131 area of about 9,000 km2. Its depositional
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