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The Devonian-Cretaceous Fossil Record of “Conchostracans” of Africa and T Their Paleobiogeographic Relationships with Other Gondwanan Faunas ∗ Oscar F

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The Devonian-Cretaceous Fossil Record of “Conchostracans” of Africa and T Their Paleobiogeographic Relationships with Other Gondwanan Faunas ∗ Oscar F Journal of African Earth Sciences 161 (2020) 103648 Contents lists available at ScienceDirect Journal of African Earth Sciences journal homepage: www.elsevier.com/locate/jafrearsci The Devonian-Cretaceous fossil record of “conchostracans” of Africa and T their paleobiogeographic relationships with other Gondwanan faunas ∗ Oscar F. Gallegoa,b, , Mateo D. Monferrana,b, Alycia L. Stigallc, Iracema A. Zacaríasa,b, Thomas A. Hegnad, Victoria C. Jiméneza, Jonathas S. Bittencourte, Gang Lif, Hugo G. Barrios Calathakia a Centro de Ecología Aplicada del Litoral, CECOAL-CONICET-UNNE, Provincial Route N° 5, Corrientes, 3400, Argentina b Geología Histórica-Paleoinvertebrados-Micropaleontología (Área Ciencias de la Tierra - Departamento de Biología), Facultad de Ciencias Exactas, Naturales y Agrimensura (FaCENA), Universidad Nacional del Nordeste (UNNE), Av. Libertad 4450, 3400, Corrientes, Argentina c Department of Geological Sciences and OHIO Center for Ecology and Evolutionary Studies, Ohio University, Athens, OH, 45701, USA d Department of Geology and Environmental Science, SUNY Fredonia, 280 Central Ave., Jewett Hall 203, Fredonia, NY 14063, USA e Laboratório de Paleontologia e Macroevolução, Centro de Pesquisas Professor Manoel Teixeira da Costa, Departamento de Geologia, Instituto de Geociências, Universidade Federal de Minas Gerais, Av. Presidente Antônio Carlos 6627, Pampulha, 31270-901, Belo Horizonte, MG, Brazil f State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, 210008, China ARTICLE INFO ABSTRACT Keywords: The main objective of this work is to present a survey of the fossil record of “conchostracans”, a group of Spinicaudata continental (also occasionally from brackish and marine sequences) fossil invertebrates, recorded from Africa, to Laevicaudata explore relationships among these taxa and those recorded from other paleocontinents, and to assess the po- Cyclestherida tential for the African “conchostracan” records to provide insight on paleobiogeographic connections within Paleozoic Gondwana. This work is focused on “conchostracans” (laevicaudatans, spinicaudatans and cyclestheriids), be- Mesozoic cause “conchostracans” are one of the most widely documented groups in the Phanerozoic continental se- Dispersal quences. A clear signal of taxonomic change is recovered within the African fauna. Leaiids, lioestheriids and palaeolimnadiids characterize African Paleozoic-Triassic strata, whereas afrograptids, fushunograptids and pa- laeolimnadiopseids characterize African Mesozoic strata. The relationship among African and other Gondwanan taxa is considered, and the potential for “conchostracans” as paleobiogeographic indicators is evaluated. Biogeographic analysis is based on the compiled data about the African “conchostacans” fossil record with specific consideration of some emblematic groups (Leaiidae, Paleolimnadidae, Vertexiinae, and Afrograptidae). Key dispersal pathways may have operated. During the Devonian to Triassic, records suggest four different faunal interchanges between Africa and Australia, Europe, South America, and India. During the Jurassic to Cretaceous, records suggest five different faunal interchanges between Africa and South America, Europe, Asia, India and Antarctica marked by different emblematic taxa. 1. Introduction authors (for more details see for insects Riek, 1973, 1976; Schlüter, 2003; mollusks by Amalitzky, 1895a, 1895b; Rossouw, 1970; ostracods Two invertebrate animal phyla, Mollusca and Arthropoda, are the by Marlière, 1950; Grekoff, 1963; Krömmelbein, 1966; Bate, 1975; main components of non-marine faunas recorded in Phanerozoic se- Colin and Dépêche, 1997; Colin et al., 1992, 1997; “conchostracans” by quences all over the world (Gray, 1988). These records have been Jones, 1862, 1878, 1890a, b, Jones and Woodward, 1894; Pruvost, known since the 19th century and are of fundamental interest for re- 1911, 1919; Leriche, 1913, 1932; Teixeira, 1943, 1947, 1958, 1960; solving biogeographical relationships among Africa and other Gond- Marlière, 1950; Kobayashi, 1954; Novozhilov, 1958b; Defretin, 1953; wana continents following the breakup of Pangea. The continental fossil Defretin et al., 1953; Defretin-Lefranc, 1967; and Tasch, 1984, 1987; invertebrate records from Africa have been studied by numerous among others see Tasch, 1987), but the group analyzed in the present ∗ Corresponding author. Centro de Ecología Aplicada del Litoral, CECOAL-CONICET-UNNE, Provincial Route N° 5, Corrientes, 3400, Argentina. E-mail addresses: [email protected] (O.F. Gallego), [email protected] (M.D. Monferran), [email protected] (A.L. Stigall), [email protected] (I.A. Zacarías), [email protected] (T.A. Hegna), [email protected] (V.C. Jiménez), [email protected] (J.S. Bittencourt), [email protected] (G. Li), [email protected] (H.G. Barrios Calathaki). https://doi.org/10.1016/j.jafrearsci.2019.103648 Received 30 December 2018; Received in revised form 3 September 2019; Accepted 16 September 2019 Available online 27 September 2019 1464-343X/ © 2019 Elsevier Ltd. All rights reserved. O.F. Gallego, et al. Journal of African Earth Sciences 161 (2020) 103648 work, “conchostracans”, was studied unevenly throughout the last two in Schwentner et al., 2012, p. 1622). In addition, Brtek and Thiéry centuries (Tasch, 1987; Anderson and Anderson, 1993; Anderson, 1999; (1995) in their study on the great European branchiopods propose that Roberts et al., 2015; Boukhalfa et al., 2017; Li et al., 2017). their distribution is related to the latitudinal gradient and the abiotic The name “conchostracan” (also known as Estheria, Estheriids, characteristics of their habitats, which also condition their diversity and phyllopods, clam shrimps, etc. see Gallego, 2010) has an extensive specific wealth. history of use in the biological and earth sciences, although that name The goal of this work is to present a survey of the continental was recently abandoned as a taxonomic unit due to the paraphyletic “conchostracans” recorded in Africa and their relationship with those relationship of its included families (Olesen, 1998). Members of the from other paleocontinents. This is done in order to evaluate the ap- “conchostracans” are branchiopod crustaceans in the Order Diplostraca plication of “conchostracan” distributions to resolve paleobiogeo- that are presently divided among the extant suborders Laevicaudata, graphic relationships within Gondwana. The broad distribution of many Spinicaudata, Cyclestherida and Cladocera and two extinct suborders species of “conchostracans” in both northern hemisphere and Leaiina and Estheriellina (Martin and Davis, 2001; Shen, 2003; Li, Gondwana during late Paleozoic and Mesozoic provides a framework 2017). The two extinct suborders have an uncertain relationship with for paleobiogeographic study. Specifically, we will examine biogeo- the rest of the “conchostracans”. In the present work, we use the term graphic patterns of the emblematic “conchostracan” groups Leaiidae, “conchostracans” following the historical and general use to collec- Palaeolimnadidae, Vertexiinae, Ulugkemiidae, Afrograptidae and tively refer to laevicaudatan, spinicaudatan, cyclestherid, leaiine, and others to evaluate their potential as paleobiogeographic indicators. estherielline crustaceans. Beginning in the Devonian, “conchostracans” are very common 2. Historical background within continental sediments from different paleoenvironments (Frank, 1988; Webb, 1979) such as estuaries, deltas, lakes and ponds, and The history of the study of spinicaudatan and other “conchostracan” brackish environments (also ocassionaly or accidentally found in records from Africa extends into the 19th Century with the pioneering marine sequences, Kobayashi, 1954). “Conchostracan” eggs are de- work of Jones (1862, 1878, 1890a, b) and Jones and Woodward (1894), siccation resistant and are distributed readily by wind (Brendonck and Pruvost, (1911, 1919), Teixeira (1943, 1947, 1958, 1960), Marlière Riddoch, 1999), in animal feces (Proctor, 1964) and tracked in mud (1950), Kobayashi (1954), Novozhilov, 1958b, Defretin (1953), adhering to animal limbs. Many fossil “conchostracan” species have Defretin et al. (1953), Defretin-Lefranc (1967) and Tasch (1984, 1987). very wide recorded transcontinental distribution. During the Paleozoic Approximately a hundred African fossil species have been described and Mesozoic, species with distinct different diagnostic features became since initial the contributions of Grey (1871), Jones (1878) and Jones dominant, and a number of these species have become biostrati- and Woodward (1894) which began the published records of African graphically useful due to their short temporal ranges and very wide “conchostracans”. Those authors described the late Permian “Estheria” geographical distribution (Kozur and Weems, 2010; Martens, 2012; greyi Jones and the Triassic species “Estheria” draperi Jones and Scholze and Schneider, 2015; Scholze et al., 2016, 2018; Schneider and Woodward from the Republic of South Africa. Scholze, 2018, etc.). However, the dispersal of this group requires ad- The most substantial work to date on African “conchostracans” was ditional future analysis since there are many questions still unanswered. Tasch's (1987) landmark monograph, “Fossil Conchostraca of the For example, what drove the evolution of the branchiopod life cycle in Southern Hemisphere and Continental Drift.” In this volume,
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