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Gymnosperms from the Lower Cretaceous Crato Formation (Brazil)

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Gymnosperms from the Lower Cretaceous Crato Formation (Brazil) Mitt. Mus. Nat.kd. Berl.. Geowiss. Reihe 7 (2004) 155-174 10. 10.2004 Gymnosperms from the Lower Cretaceous Crato Formation (Brazil). I. Araucariaceae and Lindleycladus (incertae sedis) Lutz Kunzmann', Barbara A. R. Mohr2 & Mary E. C. Bernardes-de-Oliveira3 With 1 figure and 8 plates Abstract Fossil conifers from the Early Cretaceous, most likely late Aptian, Crato Formation were studied. The excellent preservation of several of those fossils allowed detailed investigations of the leaf epidermis by light microscope (LM) and by scanning electron microscope (SEM). Members of two conifer taxa were recognized: The Araucariaceae are represented by a female cone of cf. Aruucuriu spec. A juvenile cone (Araucuriostrobus spec.) and sterile foliage shoots of Brachyphyllum obesuni might be attributed to the Araucariaceae as well. The morpho-genus Lindleycludus (incertae sedis) is represented by foliage shoots. The occurrence of Lzndleycludus is the first record of this extinct morpho-genus in the Southern Hemisphere. Anato- mical features of these conifers are interpreted as adaptations to a warm seasonally dry climate. Taphonomic problems concer- ning these conifer remains are discussed. Key words: Fossil conifers, leaf epidermis, Araucariaceae, Lindleycludus, Early Cretaceous, Crato Formation, Brazil. Zusammenfassung Fossile Koniferen aus der unterkretazischen, hochstwahrscheinlich oberaptischen Crato Formation wurden untersucht. Die ausgezeichnete Erhaltung einiger Exemplare gestattete die Analyse von Blattepidermen mittels Licht- und Rasterelektronen- mikroskop. Vertreter zweier Koniferentaxa wurden nachgewiesen: Araucariaceae kommen mit einem weiblichen Zapfen von cf. Araucuriu spec. vor. Ein juveniler Zapfen (Araucuriostrobus spec.) und sterile Zweige mit der Beblatterung der Morpho- Species Brachyphyllum obesum werden unter Vorbehalt ebenfalls zu den Araucariaceae gestellt. Vegetative Organe der Mor- pho-Gattung Lindleycludus (incertae sedis) sind ebenfalls erhalten. Das Vorkommen von Lindleycludus ist ein erster Nach- weis dieser fossilen Morpho-Gattung in der Sudhemisphare. Anatomische Merkmale dieser Koniferen werden als Anpassun- gen an ein warmes, saisonal trockenes Klima gedeutet. Im Zusammenhang mit den Koniferenresten werden taphonomische Probleme diskutiert. Schliisselworter: Fossile Koniferen, Blattepidermis, Araucariaceae, Lindleycladus, Untere Kreide, Crato Formation, Brasilien. Introduction ern Japan, Afrocedrus grows in Ethiopia and the Podocarpaceae Dacrycarpus and Dacrydium During the Early Cretaceous many conifer fa- reach SE-Asia (Page 1990). milies, such as the Araucariaceae, Podocarpaceae, Earlier studies on Mesozoic conifers were fo- Cupressaceae s. 1. (including Taxodiaceae) , and cused mostly on the Northern Hemisphere, how- Cheirolepidiaceae were present worldwide. While ever, during the last decades Southern Hemi- the Cheirolepidiaceae are now extinct, Araucaria- sphere conifers were studied intensely as well ceae and Podocarpaceae grow mainly in the (Hill & Brodribb 1999; see below). High latitude Southern Hemisphere. There are several excep- Cretaceous microfloras from the Antarctic Pe- tions e.g. Agathis extends to Maleysia and the ninsula area, the Kerguelen Plateau, and Austra- Phillipines, Nageia occurs in Asia as far as south- lia contain pollen of various Podocarpaceae s. l., Museum of Mineralogy and Geology of the State Collections of Natural History Dresden, Konigsbrucker Landstr. 159, 01 109 Dresden, Germany. E-mail: [email protected] * Museum of Natural History of the Humboldt University Berlin, Department of Paleontology, Invalidenstrasse 43, 10115 Berlin, Germany. E-mail: [email protected] Institute of Geosciences, University of Siio Paulo, Cidade Universitaria, Rua do Lago 562, SHo Paulo, CEP 05508-080, Brazil and Laboratory of Geosciences of the University of Guarulhos, Brazil. E-mail: [email protected] Received March 2004, accepted June 2004 0 2004 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 156 Kunzmann. L. et al., Gymnosperms from the Lower Cretaceous such as Podocarpus. Microcachrys, Dacrydium, The plants are thought to be mainly from terres- and Phyllocladus of the Phyllocladaceae (Cook- trial environments, but there are also a few puta- son & Pike 1953, Dettmann et al. 1992). Macro- tive aquatic elements. floras reflect the importance of conifers in the The conifer component from the Crato and southern vegetation as well. In the late Albian of Santana Formations was previously known Alexander Island various species of Araucaria- mainly from the pollen record, but also from se- ceae, Podocarpaceae and Cupressaceae s. 1.. and veral megaremains. Araucariaceae are repre- morpho-genera such as Podozamites, Elatocla- sented by pollen of the Araucariacites morpho- dus, Brachyphyllum and Pagiophylluni were species (Lima 1979) and by araucarian cone identified which make up 15% of the species di- scales (Martill et al. 1993). Lima (1979) recog- versity of this site (Cantrill & Falcon-Lang 2001). nised Podocarpaceae by its pollen. Leafy shoots From Early Cretaceous Australian strata several of Podozamites sp. (incertae sedis) which might araucarian species with various leaf types were belong to Lindleycladus (see below) are reported described (Cantrill 1992). Patagonian Mesozoic by Duarte (1985) and by Martill et al. (1993). conifer remains comprise several cones organi- Two previously described Brachyphyllum mor- cally attached to vegetative structures and include phospecies, B. obesum Heer and B. castilhoi Podocarpaceae, Cheirolepidaceae, and Araucar- Duarte were not assigned to a family due to the iaceae (e.g. Archangelsky 1994, Calder 1953, lack of diagnostic epidermal features. These taxa Stockey 1982, 1994). Floras from low latitude are also known from the Albian Santana Forma- South America are less well known (see below), tion of the same basin (Martill et al. 1993). The reports exist only from the Crato and Santana extinct family Cheirolepidiaceae is represented Formations of Brazil and from Colombia (e.g. in the flora by pollen of the genus Classopollis Pons et al. 1992). Pflug (Lima 1979) and by vegetative shoots of This study is focussed on conifer specimens the morpho-genus Pseudofrenelopsis Nathorst in with preserved organic matter and on slabs with Felix & Lenk, 1893 emend. Watson, 1977. This partly silicified plant remains from the late Ap- taxon is present in the Crato as well as in the tian Crato Formation of Brazil and thus adds Santana Formation (Bernardes-de-Oliveira, oral information on conifer taxa of Southern Hemi- comm.). Another cheirolepidiacean taxon, To- sphere low latitudes. Descriptions of the epider- maxellia Archangelsky 1963, previously de- mis anatomy of leaf taxa are given, which have scribed from Patagonia (Archangelsky 1963, been known from this flora as morpho-genera 1966, 1968) has been observed as well (present (Duarte 1985). In addition, we present prelimin- in the collection of the MfN, Berlin) and will be ary results on conifer taxa of which the repro- described in more detail in a forthcoming paper. ductive structures are preserved. In addition to those taxa cited several undeter- mined conifer remains have been mentioned from the Crato Formation, such as “small cones, The Flora of the Crato Formation incertae sedis” (Martill et al. 1993). Plant fossils and sporomorphs from the Crato Formation have been known for many decades, The fossil record of selected conifer families in and several papers on the flora have been pub- South America lished to date (Barreto et al. 2000, Duarte 1985, Maisey 1991, Lima 1978, Lima et al. 1989, Mohr Today four conifer families grow in the southern & Friis 2000, Mohr & Rydin 2002, Pons et al. hemisphere, three of them in South America: 1992). The Crato flora comprises a diverse as- Araucariaceae, Cupressaceae and Podocarpa- semblage of free-sporing plants, various gymnos- ceae. In addition, in the geologic past the extinct perms such as cycadophytes, gnetophytes (Os- Cheirolepidiaceae were present. Here, we sum- born et al. 1993, Pons et al. 1992, Rydin et al. marise briefly the fossil record of these families 2003, Mohr et al. 2004) and conifers (Duarte in South America, except for the Cupressaceae 1985) in addition to a variety of angiosperms s. 1. which seem to be irrelevant for this paper. (Mohr & Friis 2000). In total about 70-80 spe- The Araucariaceae contain three extant genera, cies have been recognized, of which more than Agathis, Araucaria and Wollemia (Farjon 1998). 20 taxa are thought to represent angiosperms, While the fossil record of Agathis and Wollemia mostly magnoliid dicotyledons (Mohr & Eklund seems to be restricted to Australia and New 2003, Mohr & Bernardes-de-Oliveira, in press). Zealand (Chambers et al. 1998, Enright & Hill Mitt. Mus. Nat.kd. Berl.. Geowiss. Reihe 7 (2004) 157 1995), Araucaria was undoubtedly present in the ceae in South America, Saxegothea, Prumnopitys Mesozoic of South America. and Dacrydium bear scale-like or awl-like leaves The section Araucaria of the genus Araucaria, and seem to be not relevant for this study. with two extant species in South America, is re- Podozamites, Lindleycladus, Angariella belong presented in the fossil record by leaf remains according to Harris (1979) to the poorly defined from the Tertiary of the Rio Negro, Argentina family Podozamitaceae which consisted of several (Menendez & Caccavari 1966, Stockey 1994). vegetative and of several fertile morpho-genera The section Bunya is restricted to a single mod- (Swedenborgia,
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