Journal of African Earth Sciences 161 (2020) 103648

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Journal of African Earth Sciences

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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, Paul general (Weeks et al., 1997), and what drove the evolution of eggs that Tasch compiled all the previous systematic works, described new spe- could pass through digestive tracts? From the Triassic onward, flying cies, and provided consistent descriptions for all known Gondwana vertebrates could have migrated and transported eggs through their “conchostracan” species. In that contribution, Tasch (1987) stated that feces (i.e. pterosaurs). Also, migrating non-volant vertebrates as dicy- “nineteen African countries are now known to possess Paleozoic-Mesozoic nodonts, laberyntodonts, temnospondils, rhynchosaurs and arch- nonmarine, conchostracan-bearing deposits”. In addition, Tasch (1987) osauriforms from northern and southern hemisphere were mentioned mentioned and analyzed some genera and species, called bioprograms by Gallego and Melchor (2000) as potential “conchostracan” eggs (e.g., Estheriina, Cornia, Palaeolimnadia, Cyclestherioides, Leaia, Pseu- transport. However, the long-distance dispersal before the evolution of doasmussiatta, Estheriella, Palaeolimnadiopsis) as important indicator the vertebrate flight and after this evolutionary event was surely han- taxa for examining biogeographic patterns within Gondwana. dled by the wind systems. Previous to this evolution step and during the Paleozoic time, wind transport is likely to be effective across long dis- 3. Material and methods tances as supported by Brendonck and Riddoch (1999) for recent forms and Gallego and Melchor (2000) for Paleozoic-Triassic and Gallego and A first database of all published Gondwana “conchostracan” oc- Rinaldi (2001) for late Mesozoic “conchostracans” both based in currences (see Table 1, Figs. 1–2 and Supplementary data) was com- southern South American records. piled based mainly on the work of Tasch (1969, 1987) combined with Dispersal of resting eggs is always passive and may be mediated by pioneering contributions from Jones (1862), Raymond (1946), wind, floods, or biotic vectors such as waterfowl (Bilton et al., 2001). In Kobayashi (1954), Novozhilov, 1958b, Defretin-Lefranc (1967), Zhang this latter case, the eggs either stick to the birds’ feathers or are carried et al. (1976) and Chen and Shen (1985) and augmented with in- internally after the ingestion of egg-bearing females. This mode of formation published in the last thirty years. This database is primarily a dispersal is not available to other aquatic taxa such as fish or mollusks literature compilation, and thus represents a springboard for this and and allows “large branchiopods” to disperse among unconnected water future analyses. Although species are provided the best taxonomic as- bodies. As aquatic species with resting eggs are generally among the signments possible based on the published information, no new ana- first colonizers of new freshwater habitats, it has been assumed thatthe lyses of type specimens, SEM work, or morphometric analyses were dispersal potential of resting eggs is high (Bilton et al., 2001, in undertaken as part of this compilation. The goal of this present work is Schwentner et al., 2012, p. 1606). In summary, the congruent pattern of to develop the first comprehensive database of published occurrences as virtually no genetic differentiation over large areas (Murray–Darling, a way to survey the diversity of “conchostracans” from Africa. Bulloo, and south and west Lake Eyre Basins, approximately In the preparation of this database, we selected every mention of 800,000 km2), a phenomenon that has not been observed to this extent “conchostracans” from the Devonian to Cretaceous of the Southern in any other branchiopod species, may be attributable to the dispersal Hemisphere. The original systematics were analyzed and then corrected capacity of the abundant, vagile migratory water birds that inhabit east according to the current systematic scheme of Chen and Shen (1985) and central Australia during wet periods. Wind and local floods may and completed with other contributions (as Zhang et al., 1976; further facilitate the dispersal of resting eggs locally (Bilton et al., 2001, Novozhilov, 1958b; Jones and Chen, 2000; Martin and Davis, 2001;

2 O.F. Gallego, et al. Journal of African Earth Sciences 161 (2020) 103648

Fig. 1. Geographical location of the chronological “conchostracan” records of the Africa continent (modified from Tasch, 1987 and compiled from different sources, see supplementary data A) a. Devonian–Carboniferous–Permian times. b. Triassic/Triassic–Jurassic times. c. Triassic–Jurassic/Jurassic/Jurassic–Cretaceous. d. Jurassic–Cretaceous/Cretaceous.

Kozur and Weems, 2010; Astrop and Hegna, 2015; Scholze and without a thorough re-evalutation including SEM imagery, many of Schneider, 2015; among others). When assignments were doubtful, we those specimens cannot be concretely placed. It is our opinion that in reviewed the original diagnosis, descriptions and figures to reach a certain cases, “conchostracan” specimens that are poorly preserved are more supported systematic assessment of the taxa. On the other hand, taxonomically useless. Also, reevaluating some of Tasch's African ma- when stratigraphic, chronologic or geographic information is incorrect terial has only reinforced this perception. or outdated, we tried to update this information when possible. In many Figs. 1–2 are made based on this database (see Supplementary data) cases, we took the original taxonomy at face value. Many of the species and when the chronological data are imprecise (as Triassic to Cretac- described in the older literature were insufficiently illustrated, and eous or others) species are placed in all graphs of the corresponding

3 O.F. Gallego, et al. Journal of African Earth Sciences 161 (2020) 103648

Fig. 2. “Conchostracan” chronological frequencies of species from African records (based in Tables 1 and 2; compiled here). ages. We have updated Tasch's (1987) nineteen records to twenty-one text mentions of "Estheria" that lack photographic documentation and included the current official name of each country (Fig. 1), which (Tasch, 1987). Nevertherless, there are very strong sampling biases now includes: People's Democratic Republic of Algeria, Republic of within continental records in general within both Afria and globally, Angola, Republic of Cameroon, Republic of Chad, Democratic Republic which impact the “conchostracan” records. of the Congo (formerly Belgian Congo or Zaire), Gabonese Republic (formerly Gabon), Republic of Ghana, Republic of Equatorial Guinea 4.1. Geographic and temporal distribution of African “conchostracans” (formerly Rio Muni or Spanish Guinea), Republic of Kenya, Kingdom of Lesotho, Republic of Madagascar, Republic of Malawi (formerly Nya- The fossil record of “conchostracans” demonstrates clear trends saland), Kingdom of Morocco, Republic of Mozambique, Republic of across geographic space and geologic time. Notably and despite the Niger, Republic of South Africa, United Republic of Tanzania, Republic sampling biases, different faunas can be identified for the Paleozoic and of Zambia (formerly Northern Rhodesia), and Republic of Zimbabwe. In Mesozoic as described in the following text. addition, we include new records from Republic of Namibia and Re- The Paleozoic Era is represented by records of Devonian to Permian public of Tunisia (Stigall et al., 2014; Boukhalfa et al., 2017; Li et al., faunas. The Devonian includes only one dubious record of a probable 2017). estheriteoidean or “lioestheriid” (“Lioestheria” sp. Rennie, 1934). Occurrences of Africa “conchostracan” species are compiled from African Paleozoic “conchostracan” records show a large number of various published sources as well as the authors' own data. The pre- leaiid species, mainly in the Permian Period, with a dominance of the sence of “conchostracans” in Africa and their relationships with the genus Hemicycloleaia. Euestheriids have also been recorded in Paleozoic paleobiogeography is analyzed here focused in two main points: one sequences with five species cited by Tasch (1987, see Table 1 and related to individual species (like Euestheria minuta), and the other Supplementary data). Also, species of the families Orthothemosiidae (2 genus (like Congestheriella) or family levels with records outside the spp.), Estheriellidae (2 spp.), Lioestheriidae (3 spp.) and Esther- African continents and the temporal distribution and Gondwanan pa- iteoidean indet (2 spp.). are known from the late Carboniferous leogeography. (Termier and Termier, 1950; Bonnet et al., 1960; Tasch, 1987); how- The following abbreviations are used across this contribution: USTL ever, they are in need of systematic revision. Nevertheless, there are (Lille University – Sciences and Technologies; because, as formerly around six indeterminate spinicaudatan species of Paleozoic age (see called Université des Sciences et Technologies de Lille, France, its ac- Table 1, Fig. 2). ronym was USTL); MGL, Musée Gosselet, Lille, France (Musée The African “conchostracan” fauna is relatively stable from the late d’Histoire Naturelle– Musée de Géologie) and USNM, United States Carboniferous through Triassic. The lioestheriids (Vertexinae) plus two National Museum (Smithsonian), Washington, D.C. estheriellids, one ulugkemiid and one estheriteoidean characterized the late Carboniferous, early Permian and Early and Late Triassic times. 4. Results Also, the great record of euestheriids and the genus Euestheria (the most diverse group) characterize this time span (late Carboniferous, late The greatest concentration of “conchostracans” bearing deposits Permian, and Early and Late Triassic) in Africa. The faunal similarities occurs south of the equator, as it is the case with South America between the late Carboniferous to Late Triassic are only marked by the (Gallego, 2001a, b; Gallego and Martins-Neto, 2006). Democratic Re- disappearance of the leaiids at the end-Permian extinction, however public of Congo (DRC) has the greatest concentration of fossiliferous these results are based on a low number of recorded species. sites (see Fig. 1). Angola has the next highest concentration, followed The Mesozoic “conchostracans” of Africa, particularly the Triassic by the Republic of South Africa (RSA), Zimbabwe, and Lesotho (see faunas, exhibit higher diversity and abundance than the Paleozoic Fig. 1). North of the equator, Gabon, Niger, Algeria, and Morocco each faunas despite the above mentioned sampling biases. The post- have several sites. From most of the remaining countries published Paleozoic-Triassic interval starts with three records of Eosestheriidae, records indicate only a single species (ex. Stigall et al., 2014) or are in- Fushunograptidae and indeterminate species during the Early Jurassic.

4 O.F. Gallego, et al. Journal of African Earth Sciences 161 (2020) 103648

Table 1 Records of Paleozoic “conchostracan” species of Africa according to age/stratigraphic unit (for more information see Supplementary Data).

PALEOZOIC

Family Genus Species Age/Stratigraphic unit

Euestheriidae Euestheria anchietai (Teixeira, 1947) Marlière, late Permian/Lutoe fish beds - Cassanje I. 1950 Euestheriidae Euestheria greyi (Jones, 1878) Tasch, 1987 late Permian - early Triassic/Shales fish beds, upper Maji ya Chami beds - Middle Beaufort/Dark grey shale near Cradock, Cape Province. Euestheriidae Euestheria cf. mangaliensis (Jones, 1862) Bond, late Permian/Upper Madumabisa Shales. 1955 Orthothemosiidae Pseudestheria welleri late Permian or Early Triassic/Escarpment Grit (near base of Upper Karoo) Bond, 1964 (in Tasch, 1987) Orthothemosiidae Pseudestheria sp. late Permian - Upper Madumabisa Shales - Lower Karoo. Bond (1952) Estheriteoidean indet "Lioestheria" borgesi late Permian/Karoo Formation - Lower Shire-Zambesi region, Plant bearing beds. Teixeira (1943) Family indet. Isaura cf. mangaliensis (see Supl. Data) late Permian/Upper Matabola Beds (Jones, 1862) Bond, 1955 Family indet. Cyzicus sp. Permian-Triassic/Mafungabusi Plateau - Forest Sandstone Haughton (1927) Family indet. "Estheria" sp. late Permian/Lower and Middle Madumabisa Shales. Lightfoot (1914) Family indet. "Estheria" sp. late Permian - Sakamena facies. Besairie (1952) Family indet. "Estheria" (C.) sp. Lower Beaufort - uppermost Ecca/Red marls with fish scales. Haughton (1963) Family indet. Cyzicus(E.) sp. early Permian - Middle Ecca. Plant bearing Roux (1960) Estheriellidae Nyasaestheriella nyasana (Newton, 1910) Permian? - Karoo beds, Nkana Kobayashi, 1954 Leaiidae Hemicycloleaia cf. normalis late Permian - Middle Madumabisa Shales - Lower Karoo. Bond (1952) Leaiidae Hemicycloleaia cf. haynsi late Permian - Upper Madumabisa Shales - Lower Karoo. Bond (1952) Leaiidae Hemicycloleaia cradockensis late Permian - Upper Karoo Series - Middle Beaufort. Tasch (1987) Leaiidae Hemicycloleaia sengwensis late Permian - Upper Madumabisa Mudstone Bond, 1955 (in Tasch, 1987) Leaiidae Hemicycloleaia sessami late Permian - Middle Madumabisa Shales - Lower Beaufort. Bond, 1955 (in Tasch, 1987) Leaiidae Hemicycloleaia sp. late Permian - Lower Beaufort Beds. Barnard, 1931 (in Tasch, 1987)

Lioestheriidae Gabonestheria gabonensis (Marlière, 1950) early Permian/Late Triassic - Red mudstones, Agoula River Series/Cassanje III Series Phyllopod Novozhilov, 1958b Beds Stage.

Lioestheriidae Cornia? limbata (Goldenberg, 1877) Kobayashi, Carboniferous-Lower Westphalian/Columb Bechar-Kenadze Basin. 1954 Estheriellidae Estheriella? sp. late Carboniferous - Westphalian/Hamahou Ida ou Zal. Termier and Termier, 1950 (in Tasch, 1987) Euestheriidae Euestheria simoni (Pruvost, 1911) Pruvost, 1919 Carboniferous-Lower Westphalian/Kenadze Basin - Sidi Bel Hassam-Djerada-Sidi Bel Kessem sondage. Euestheriidae Euestheria subsimoni (Deleau, 1945) Tasch, 1987 Carboniferous-Lower Westphalian/Columb Bechar-Kenadze Basin. Leaiidae Leaia bertrandi Carboniferous - Lower Stephanian/Haute Atlas-Hamalou-Agadir. Tasch (1987) Leaiidae Hemicycloleaia sp. 1 Carboniferous - Westphalian/Hamahou Ida ou Zal. Termier and Termier, 1950 (in Tasch, 1987) Leaiidae Hemicycloleaia sp. 2 Carboniferous - Westphalian/Hamahou Ida ou Zal. Termier and Termier, 1950 (in Tasch, 1987) Rostroleaiidae Rostroleaia sp. Carboniferous - Namurian? to Stephanian? - Reggan Basin Bonnet et al. (1960)

Estheriteoidean indet "Lioestheria"? sp. early Devonian/Witteberg Series Cape System. Rennie, 1934 (in Tasch, 1987)

Paleolimnadiidae and Euestheriidae are well represented with many Triassic and Early Jurassic by the appearance of the families Afrograptidae, species belonging to the genera Palaeolimnadia and Euestheria. In ad- Cyclestheriidae and Sinoestheriidae and the possible disappearance of the dition, new families such as Asmussiidae, Aquilonoglyptidae, families Aquilonoglyptidae, Palaeolimnadiidae, Perilimnadiidae and Perilimnadiidae and Paleolynceidae appear in the Triassic, and other Paleolynceidae. Most spinicaudatans (e.g., Palaeolimnadiidae and families like Estheriellidae and Lioestheriidae appear after the Euestheriidae) and Estheriellidae show a marked diversity increase during Paleozoic. Following this, diversity declines to the Jurassic level with a Triassic and then decline in Jurassic. In contrast to these families, drop in taxonomic richness probably due to collection biases and pos- Fushunograptidae and Afrograptidae record diversity increase during sible loss of records. Jurassic and Cretaceous. The estheriellids characterize the Late Jurassic in The character of the “conchostracan” fauna changes between the Africa. Also, other families like Cyclestheriidae, Sinoestheriidae,

5 O.F. Gallego, et al. Journal of African Earth Sciences 161 (2020) 103648

Palaeolimnadiopseidae, Lioestheriidae, Euestheriidae, Orthothemosiidae Fig. 3B; Tasch, 1987; Shen, 2006). However, palaeolimnadids are rare and Estheriellidae are known from the Upper Jurassic units, but more in South America; they only include Palaeolimnadia herbsti from Chile specifically by Fushunograptidae. Furthermore, species of Sinoestheriidae (Upper Permian, Gallego and Breitkreuz, 1994) and Palaeolimnadia sp. and Fushunograptidae are known from the Late Jurassic-Early Cretaceous from Brazil (Upper Jurassic-Lower Cretaceous; Tasch, 1987). On the (or simply Jurassic-Cretaceous as reported by Roberts et al., 2015). The other hand, palaeolimnadids Estheriina and Palaeolimnadia are known Cretaceous has high-diversity records with eleven families that show a low from India and Antarctica (Lower Jurassic, Tasch, 1987). With respect abundance, and some families (Loxomegaglyptidae and Anthro- to the Northern Hemisphere records (Central Europe, Asia and North nestheriidae) appear only in this period (see Fig. 2, Table 2). America), Palaeolimnadia occurs in the Triassic (Kozur and Weems, Families Paleolimnadidae, Lioestheriidae, Euestheriidae, Eosestheriidae, 2010). Asmussiidae, and Afrograptidae are known from the Early Cretaceous time, The Family Palaeolimnadiopseidae (Superfamily Vertexioidea, see which probably has a higher diversity and is best represented considering Astrop and Hegna, 2015) has three genera and five species recorded the number of stratigraphic units and sampling biases. The Family from the Mesozoic of Congo and Angola: Palaeolimnadiopsis, Pterio- Fushunograptidae also characterizes this stage (as previously mentioned for grapta and the endemic Stanleyviella (Teixeira, 1958; Defretin-Lefranc, the Upper Jurassic units). Finally, the families Euestheriidae, 1967; Tasch, 1987). Pteriograpta cf. reali is also known from South Loxomegaglyptidae and Orthothemosiidae are known from the Late America (Cardoso, 1971; Tinoco and Katoo, 1975; Tasch, 1987, actively Cretaceous span that is also characterized by Antronestheriidae, being studied by one of us, JSB) and Macrolimnadiopsis from India Fushunograptidae and Afrograptidae. (Ghosh, 2011) and South America (Beurlen, 1954). Nevertheless, the Some stratigraphic units with ambiguous or dubious ages (Haute African diversity is low and similar to that of Australia with two Triassic Lueki Series, Cassanje III, see Table 2, Supplementary data) for example species records. It is likewise similar to the diversity of species from Late Triassic or Jurassic (pre-Oxfordian) units compiled from Tasch India with six known Permian-Triassic species (Tasch, 1987; Ghosh, (1987) are composed by faunas that include Palaeolimnadiopseidae, 2011). This is in stark contrast to South America, where seventeen Lioestheriidae, Euestheriidae, Eosestheriidae, Orthothemosiidae, Afro- species are recorded, ranging from the late Permian to the Early Cre- graptidae and Estheriellidae. In these cases, comparisons made with taceous (Rohn and Cavalheiro, 1996). other southern faunas suggest a Triassic age based on records of Es- The Family Lioestheriidae (sensu Kozur et al., 1981) is known from theriella moutai and Euestheria mangaliensis reported for the Cassanje III the Carboniferous to the Lower Cretaceous sequences in Africa by true from Angola. On the other hand, the presence of Congestheriella luala- (umbonal node or spined) genera belonging to Subfamily Vertexiinae: bensis, Carapacestheria malangensis and Pteriograpta reali suggest a Jur- Echinestheria, 1 sp., Cornia 4 spp., Gabonestheria 1 sp. (Marlière, 1950; assic to Early Cretaceous age as mentioned by Roberts et al. (2015) Kobayashi, 1954; Novozhilov, 1958b; Tasch, 1984, 1987) and also by from the Lualaba Series. the related Family Perilimnadiidae Falsisca, 1 sp.; Shen, 2006). Also, the dubiously valid genus Palaeestheria (3 spp. with a large smooth um- 4.2. “Conchostracan” faunal relationship between Africa and other bonal area) is known from the Upper Jurassic to Lower Cretaceous in continents Congo and South Africa (Barnard, 1931; Defretin-Lefranc, 1967). The Permian-Triassic genus Cornia is widely known with species from South In this section, we compare “conchostracans” from Africa with other America (4 spp., Tasch, 1987; Gallego and Breitkreuz, 1994; Tassi et al., records from Gondwana and northern hemisphere continents to estab- 2013; Tassi, 2016) and Australia (2 spp.; Tasch and Jones, 1979; Tasch, lish paleobiogeographic distribution patterns. The information was 1987). It is also known from the Lower Jurassic of Antarctica (2 spp.; compiled in systematic order. Tasch, 1987). On the other hand, the lioestheriids are diverse in the The Suborder Cyclestherida (and Family Cyclestheriidae) is a Lower Triassic of India and contain the following genera that are known monogeneric taxon that contains the genus Cyclestherioides and is only from both Africa and India: Echinestheria, Cornia and Gabonestheria with known from scarce records. The genus Cyclestherioides is known from seven species. Genera Vertexia (1 sp.), Protomonocarina (2 spp.) and the Middle and Upper Permian of South America (2 spp., Tasch, 1987; Curvacornutus (4 spp.) are known in India and also from the Upper Gallego and Breitkreuz, 1994) and from the Lower Jurassic of Antarc- Paleozoic and Triassic of North America and Europe. Endemic forms tica (2 spp., Tasch, 1987). In contrast, it is also known from the Upper Turricornia (1 sp.) and Indomonocarina (1 sp.) are also recorded in the Jurassic of Tanzania of Africa (Kobayashi, 1954; Tasch, 1987). The Lower Triassic from India (Ghosh, 2011). other non-spinicaudatan record corresponds to the discovery of the The Family Ulugkemiidae (Superfamily Eosestherioidea) contains Suborder Laevicaudata (including the Paleolynceioidea). The genus the genus Kenyaestheria that is known from Lower Triassic (Kenya) in Prolynceus is known from the Lower Triassic of Kenya, which represents Africa (Shen, 2006). Ulugkemiids (by the genus Triasulugkemia) are also the only record of this small group from the Southern Hemisphere known from Middle-Upper Triassic in South America (Gallego and (Shen, 2006). Melchor, 2000). Previously, Novozhilov, 1958b defined this family With regard to the Suborder Spinicaudata, the Superfamily based on northern hemisphere records, which contains the genera: Vertexioidea contains the following five families that are known from Ulugkemia (with doubtfull validity) from the Middle Devonian and Africa: Paleolimnadidae, Paleolimnadiopseidae, Sinoestheriidae, Upper Permian and Beligum from Upper Devonian, both from Russia. Lioestheriidae and Perilimnadiidae. The endemic genus Afrolimnadia is Genera Tjulbaria and Elegestia are known from the Middle Devonian, known from the Upper Triassic in South Africa (Tasch, 1984, Table 2). and Vjatkium and Dneprium from the Upper Permian in Russia The genus Palaeolimnadia (with 10 spp.) is known from Kenya, Angola (Novozhilov and Varentsov, 1956; Novozhilov, 1959, 1961). Genera and Lesotho form the Lower and Upper Triassic of Africa (Tasch, 1984, Tshuvashium (upper Permian, Tchouvachie-Russia), Rohdendorfium 1987; Shen, 2006). The genus Estheriina (2 sp.) is known from Angola (Permian-Triassic, Rusia) and Defretinia (Upper Jurassic, Transbaikalia- (Late Triassic, Tasch, 1987) and Niger (Early Cretaceous, Tasch, 1987). Russia) probably belong to the Family Ulugkemiidae (as suggested by African palaeolimnadiids share Palaeolimnadia cf. pusilla (Shen, 2006) Gallego and Melchor, 2000). with the Lower Triassic of China (and Europe if Palaeolimnadia pusilla is The Family Euestheriidae is one of the best represented taxa in the synonym to Cornia germari as proposed by Kozur and Weems, 2010). African records, with twenty-one species from the Upper Carboniferous The genera Palaeolimnadia (11 spp., Tasch, 1987) and Estheriina (3 to the Upper Cretaceous of Angola, Algeria, Cameroon, Congo, Kenya, spp., Tasch, 1987) are also known from the Upper Permian and Upper Lesotho, Madagascar and Zimbabwe. This family contains the following Triassic in Australia and exhibit the same high diversity and also share genera that are known from Africa: Euestheria and Pseudoasmussia? species with Africa (e.g., Palaeolimnadia banksi, Palaeolimnadia cf. wia- (Raymond, 1946; Defretin-Lefranc, 1967; Tasch, 1984, 1987; Shen namattensis Fig. 3A, Palaeolimnadia cf. glabra, Estheriina cf. rewanensis et al., 2002; Shen, 2006). The genus Euestheria is the most common

6 O.F. Gallego, et al. Journal of African Earth Sciences 161 (2020) 103648

Table 2 Records of Mesozoic “conchostracan” species of Africa according to age/stratigraphic unit (for more information see Supplementary Data).

MESOZOIC

Family Genus Species Age/Stratigraphic unit

Euestheriidae Euestheria lefranci Tasch, 1987 Late Cretaceous/Ain-el-Hadjadj - Plateau du Tademait. Euestheriidae Euestheria lerichei Marlière, 1950 (in Tasch, 1987) Late Cretaceous/Kwango Group - Kabinda area - Upper Triassic/Cassanje III. Loxomegaglyptidae Gen. Indet. sp. Roberts et al., 2015 Late Cretaceous/Kwango Group - Kabinda area. Loxomegaglyptidae Gen. Indet. sp. Shen et al., 2002 Late Cretaceous. Anthronestheriidae Ethmosestheria mahajangaensis Stigall and Hartmann, Late Cretaceous/Mahajanga Basin- Berivotra Formation. 2008 Anthronestheriidae Pseudestherites sp. Roberts et al., 2015 Late Cretaceous/Kwango Group - Mbuji-Mayi/Kabinda area. Family indet. "Estheria" sp. Haughton, 1963 Late Cretaceous/Léré Series. Afrograptidae Congestheriella sp. Roberts et al., 2015 Late Cretaceous/Kwango Group - Kabinda area.

Palaeolimnadiidae Estheriina (N.) marginata (Defretin et al., 1956) Tasch, Early Cretaceous/Kori Ezazel Agades 1987 Lioestheriidae Palaeestheria sp. Barnard, 1931 Early Cretaceous/Port Elizabeth. Lioestheriidae Palaeestheria anomala (Jones, 1901) Barnard, 1931 Early Cretaceous/Uitenhage Series. Euestheriidae Euestheria cf. dahurica (Chernyshev, 1930) Defretin, Early Cretaceous/Mayo Tafal. 1953 Tasch, 1987 Euestheriidae Euestheria sambaensis Defretin-Lefranc, 1967 Early Cretaceous (post Wealden)/Kisamu Nzambe, Bokungu Series. Euestheriidae Pseudoasmussia? banduensis Defretin-Lefranc, 1967 Early Cretaceous/Loia Series. Euestheriidae Pseudoasmussia? ndekeensis Defretin-Lefranc, 1967 Early Cretaceous/Loia Series. Eosestheriidae Nigerestheria lamberti (Defretin et al., 1956) Chen (in Early Cretaceous/Kori Ezazel Agades - Pont du Mayo Louti. Zhang et al., 1976) Asmussiidae Asmussia dekeseensis Defretin-Lefranc, 1967 Early Cretaceous/Bokunga Series. Asmussiidae Asmussia ubangiensis Defretin-Lefranc, 1967 Early Cretaceous/Bokunga Series. Fushunograptidae Ordosestheria chottsensis Li et al., 2017 Early Cretaceous - Lower Barremian/uppermost Bouhedma Formation Fushunograptidae Cratostracus? tunisiaensis Boukhalfa et al., 2017 Early Cretaceous - Barremian/Sidi Aich Formation Fushunograptidae Orthestheria (Migransia) kasaiensis Marlière, 1950 (in Early Cretaceous - Wealden/Pushaluenda (Kasai) drain, Loia Series. Shen et al., 2004) Fushunograptidae "Bairdestheria" mawsoni (Jones, 1897) Raymond, 1946 Early Cretaceous/Pont du Mayo Louti. Family indet. "Estheria" sp. Greigert and Pougnet, 1967 Early Cretaceous/Irhazer Fm. Red lacustrine mudstone - Dinosaur beds Damergou. Family indet. "Estheria" sp. Furon, 1950 Early Cretaceous/Mamfe region. Afrograptidae Afrograpta tricostata (Defretin, 1953) Novojilov, 1957 Early Cretaceous/Pont du Mayo Louti. Afrograptidae Camerunograpta camerouni (Defretin, 1953) Novojilov, Early Cretaceous/Pont du Mayo Louti. 1957

Sinoestheriidae Stanleyviella sp. Roberts et al., 2015 Jurassic-Cretaceous/Stanleyville Formation - Mbuji-Mayi area. Family indet. "Estheria" sp. Haughton, 1963 Late Jurassic-Early Cretaceous/Cocobeach Series. Cyclestheriidae Cyclestherioides (C.) janenschi Kobayashi, 1954 (Tasch, Late Jurassic/Tendaguru Series. 1987) Palaeolimnadiopseidae Palaeolimnadiopsis sp. Tasch, 1987 Late Jurassic/Stanleyville Series. Sinoestheriidae Stanleyviella lombardi (Defretin-Lefranc, 1967) Chen Late Jurassic/Stanleyville Series. and Shen, 1985 Lioestheriidae Palaeestheria passaui (Marlière, 1950) Defretin-Lefranc, Late Jurassic/Stanleyville Series. 1967 Euestheriidae Pseudoasmussia? duboisi (Marlière) Defretin-Lefranc, Late Jurassic/Stanleyville Series. 1967 Fushunograptidae Australestheria corneti (Marlière) Defretin-Lefranc, Late Jurassic/Stanleyville Series. 1967, Zhang et al., 1976 Fushunograptidae "Bairdestheria" sp. Tasch, 1987 Late Jurassic-Early Cretaceous/Amisa River. Fushunograptidae Orthestheria (Migransia) biaroensis Defretin-Lefranc, Late Jurassic-Kimmeridgian/Stanleyville Series. 1967 (in Shen et al., 2004) Fushunograptidae Orthestheria (Migransia) caheni Defretin-Lefranc, 1967 Late Jurassic Kimmeridgian/Stanleyville Series. (in Shen et al., 2004) Estheriellidae Estheriella evrardi Defretin-Lefranc, 1967 Late Jurassic Kimmeridgian/Stanleyville Series.

Eosestheriidae Hardapestheria maxwelli Stigall et al., 2014 Early Jurassic/Kalkrand Fm. Fushunograptidae "Lioestheria" lebombensis Rennie, 1937 (in Tasch, 1987) Early Jurassic/Lebombo Volcanic Fm. Family indet. "Estheria" sp. Haughton, 1963 Early Jurassic/Cocobeach Series.

Palaeolimnadiidae Palaeolimnadia (P.) cf. wianamattensis (Mitchell, 1927) Late Triassic/Cassanje I Series Phyllopod Beds Stage Tasch, 1987 Palaeolimnadiidae Palaeolimnadia (G.) oesterleni Tasch, 1987 Late Triassic/Cassanje I Series Phyllopod Beds Stage Palaeolimnadiidae Palaeolimnadia (G.) africania Tasch, 1987 Late Triassic/Cassanje I Series Phyllopod Beds Stage Palaeolimnadiidae Palaeolimnadia (G.) sp. Tasch, 1984 Late Triassic/Cave Sandstone Palaeolimnadiidae Estheriina (N.) cf. rewanensis Tasch (Tasch and Jones, Late Triassic/Cassanje I Series Phyllopod Beds Stage 1979) Palaeolimnadiidae Afrolimnadia sibiriensis Tasch, 1987 Late Triassic/Cave Sandstone Palaeolimnadiopseidae Pteriograpta reali Teixeira, 1958 Late Triassic or Jurassic/Cassanje III Palaeolimnadiopseidae Palaeolimnadiopsis lubefuensis Defretin-Lefranc, 1967 Late Triassic or Jurassic (pre Oxfordian)/Haute Lueki Series Lioestheriidae Echinestheria marimbensis Marlière, 1950 Late Triassic or Jurassic (pre Oxfordian)/Haute Lueki Series - Upper Triassic/Cassanje III. Lioestheriidae Cornia angolata Tasch, 1987 Late Triassic/Cassanje I Series Phyllopod Beds Stage Lioestheriidae Cornia haughtoni Tasch, 1984 Late Triassic/Cave Sandstone (continued on next page)

7 O.F. Gallego, et al. Journal of African Earth Sciences 161 (2020) 103648

Table 2 (continued)

MESOZOIC

Family Genus Species Age/Stratigraphic unit

Euestheriidae Euestheria cf. minuta (von Zieten) Raymond, 1946 Late Triassic/Forest Sandstone/Mafungabusi (Bond, 1952) Euestheriidae Euestheria forbesi (Jones, 1862) Raymond, 1946 Late Triassic/north Constantine, Sid-Al-bou-Krizi Euestheriidae Euestheria stockleyi Tasch, 1984 Late Triassic/Cave Sandstone Euestheriidae Euestheria thabaningensis Tasch, 1987 Late Triassic/Cave Sandstone ? Euestheriidae "Estheria" destombesi (Defretin, 1950) Cuvelier et al., Late Triassic/Region d'Argana-Bigoudine, Haute Atlas. 2015 Orthothemosiidae Glyptoasmussia luekiensis Defretin-Lefranc, 1967 Late Triassic or Jurassic (pre Oxfordian)/Haute Lueki Series. Orthothemosiidae Pseudestheria lepersonnei Defretin-Lefranc, 1967 Late Triassic - Late Cretaceous/Cassanje III - Kwango Series - l'Inzia beds. Orthothemosiidae Orthothemos draperi (Jones and Woodward, 1894) Late Triassic/Shale overlying Cave Sandstone Raymond, 1946 Eosestheriidae Carapacestheria malangensis (Marlière, 1950) Everman, Late Triassic or Jurassic (pre-Oxfordian) Haute Lueki Series/Cassanje III. 2007 Asmussiidae Asmussia loockii Tasch, 1984 Late Triassic/Cave Sandstone Estheriteoidean indet "Lioestheria" lesothoensis Tasch, 1984 Late Triassic/Cave Sandstone Fushunograptidae "Lioestheria" tendagurensis Janensch, 1933 (in Tasch, Late Jurassic/Tendaguru Series. 1987) Estheriteoidean indet "Lioestheria" cassambensis Teixeira, 1960 Late Triassic ?/Cassanje III ? Fushunograptidae "Bairdestheria" kitariensis Marlière, 1950 (in Shen et al., Late Triassic or Jurassic - Late Cretaceous/Cassanje III - Lubilash System - Kwango Series, 2004) Canyon d l'Inzia. Afrograptidae Congestheriella lualabensis (Leriche, 1913) Kobayashi, Late Triassic - Jurassic/Lualaluba beds - Late Jurassic/Stanleyville Series. Jurassic - Early 1954 Cretaceous Lualaba series (Roberts et al., 2015 Cahen and Lepersonne) Estheriellidae Estheriella moutai (Leriche, 1932) Defretin-Lefranc, Late Triassic or Jurassic (pre Oxfordian)/Haute Lueki Series - Upper Triassic/Cassanje III. 1967 Estheriellidae Estheriella bornhardti Janensch, 1925 (in Tasch, 1987) Late Triassic/Hatambulo Beds, Beaufort Series.

Paleolynceidae Prolynceus sp. Shen, 2006 Early Triassic/Upper Maji ya Chumvi Formation. Palaeolimnadiidae Palaeolimnadia (P.) banksi (Tasch, 1975) Shen, 2006 Early Triassic/Upper Maji ya Chumvi Formation. Palaeolimnadiidae Palaeolimnadia (P.) cf. glabra (Mitchell, 1927) Tasch, Early Triassic/Upper Maji ya Chumvi Formation. 1987 Palaeolimnadiidae Palaeolimnadia (P.) cf. pusilla Shen (in Zhang et al., Early Triassic/Upper Maji ya Chumvi Formation. 1976) Shen, 2006 Palaeolimnadiidae Palaeolimnadia (G.) sp. Shen, 2006 Early Triassic/Upper Maji ya Chumvi Formation. Lioestheriidae Cornia? panchetella Tasch, 1987 Early Triassic/Upper Maji ya Chumvi Formation. Perilimnadiidae Falsisca sp. Shen, 2006 Early Triassic/Upper Maji ya Chumvi Formation. Ulugkemiidae Kenyaestheria gracilis Shen, 2006 Early Triassic/Upper Maji ya Chumvi Formation. Euestheriidae Euestheria mangaliensis (Jones) Raymond, 1946 Early Triassic/Upper Maji ya Chumvi Formation. Euestheriidae Euestheria minuta (von Zieten) Raymond, 1946 Early Triassic - Late Triassic/Upper Maji ya Chumvi Formation - north Constantine, Sid-Al-bou- Krizi - Region d'Argana-Bigoudine, Haute. Atlas. Euestheriidae Euestheria triassibrevis Tasch, 1987 (in Shen, 2006) Early Triassic/Upper Maji ya Chumvi Formation. Euestheriidae Magniestheria truempyi (Kozur and Seidel, 1983) Shen Early Triassic - Lower Sakamena Bed 4/Ambilobé area. et al., 2002 Orthothemosiidae Orthothemos sp. Shen, 2006 Early Triassic/Upper Maji ya Chumvi Formation. Orthothemosiidae Glyptoasmussia sp. 1 Shen, 2006 Early Triassic/Upper Maji ya Chumvi Formation. Orthothemosiidae Glyptoasmussia sp. 2 Shen, 2006 Early Triassic/Upper Maji ya Chumvi Formation. Orthothemosiidae Glyptoasmussia sp. 3 Shen, 2006 Early Triassic/Upper Maji ya Chumvi Formation. Aquilonoglyptidae Aquilonoglypta sp. Shen, 2006 Early Triassic/Upper Maji ya Chumvi Formation. Estheriellidae Estheriella? sp. Dietrich, 1939 (in Tasch, 1987) Triassic/Nord Adamawa Estheriellidae Estheriella? sp. Maufe, 1908 (Miller, 1952) Early Triassic/Upper Maji ya Chumvi Formation. euestheriid genus in Africa (17 spp.), as in other southern hemisphere America as E. mangaliensis an considered doubtfull by Tasch (1987) and records (Gallego, 1992, 2001a, b), with two late Carboniferous, three Kozur and Weems (2010). Also E. forbesi are known from the Middle- late Permian, eight Early-Late Triassic and four Early-Late Cretaceous Upper Triassic from South America (Argentina, Brasil, Gallego, 1992, species. The genus Pseudoasmussia? (included in the Family Eu- Gallego, 1996) and Upper Triassic of Groenland (Defretin-Lefranc, estheriidae sensu Hennion et al., 2015; Cuvelier et al., 2015) with three 1969). Kozur (1982) suggested that Euestheria forbesi (Jones) belongs to Late Jurassic-Early Cretaceous species is also known in Africa (Defretin- the genus Laxitextella, but research carried out by the author (OFG) to Lefranc, 1967). Euestheria minuta von Zieten, E. mangaliensis Jones and date does not agree with such a proposal, given that the ornamentation E. forbesi Jones are known from the Upper Triassic in Algeria and An- in E. forbesi is composed of small rounded areolae (Gallego, 1992), gola (Defretin et al., 1953). Nevertheless, the original location of E. though more studies are needed to clarify this species’ affinities. Other minuta came from the Middle Triassic of Europe also recorded from Asia recent SEM studies carried out by Tassi et al. (2015) and Tassi (2016, (Germany to China) and Canada (Kozur and Weems, 2010) and Middle - see Lám. 7D-E) do not support the proposal from Kozur (1982). Upper Triassic sequences in South America (Gallego, 1992, 2001a, b) Some African euestheriid species are distributed from more than (for more information on the controversial taxonomic range of Eu- one geographic location like the late Carboniferous E. simoni (Algeria- estheria minuta see Morton et al., 2017 and Sell, 2018). Moreover, the Morocco, and also occurs in late Carboniferous deposits of the second mentioned species is known as Magniestheria mangaliensis (sensu Westphalian C in Europe, see Pseudestheria simoni in Schneider and Kozur and Seidel, 1983) originally described from India (Jones, 1862) Scholze, 2018), late Permian-Early Triassic E. greyi (Kenya-South and also known from the Lower Triassic from Europe (Germany, Russia, Africa), the Early-Late Triassic E. minuta (Kenya, Algeria-Morocco, Hungary) and Asia (China) (Kozur and Seidel, 1983; Kozur and Weems, Zimbabwe) and the Late Cretaceous E. lerichei (Congo-Angola) (see 2010; Scholze et al., 2016) and Middle-Upper Triassic from South Table 2). African Late Cretaceous records finalize with the register of

8 O.F. Gallego, et al. Journal of African Earth Sciences 161 (2020) 103648

Fig. 3. Some selected taxa of “conchostracan” species from Africa. A. Palaeolimnadia wianamattensis (Mitchell), USNM PAL 424944, B. Estheriina (Nudusia) rewanensis Tasch, USNM PAL 426138, C. Cornia angolata Tasch, USNM PAL 424952, (Fig. 3A, B and C came from Oesterlen collection, northern Angola, Series, Phyllopod Beds, Upper Stage, Cassanje III, Late Triassic, Tasch, 1987), D. Cornia? limbata (Goldenberg), MGL 5099a (“Bowette” of Hirschbach at 766.50 m, Saarland, Germany, Sarrelouis Formation, Carboniferous, Lower Stephanian, see Hennion et al., 2015), E. Pseudestheria simoni (Pruvost), MGL 6031a (Roof of the coal seam Marie, Pit 9, Concession of Courrières, Group of Hénin-Liétard, Pas-de-Calais, France, see Hennion et al., 2015), F. “Estheria” destombesi (Defretin), USTL 1090c, (Argana-Bi- goudine area, western High-Atlas, Morocco, Aït Khtab Group, (Aït Khtab is now written Ait Khattab, Triassic, see Cuvelier et al., 2015). Scale bar = 1 mm. the Family Loxomegaglyptidae (genus indet. 2 spp. Roberts et al., the Upper Jurassic to Lower Cretaceous of China. African records of the 2015). Family Eoesetheriidae include the Early Jurassic Hardapestheria max- The Family Orthothemosiidae is a controversial taxon (considered welli (Stigall et al., 2014), Carapacestheria malangensis (Late Triassic- along with Aquilonoglyptidae as part of the Eosestherioidea as sug- Early Jurassic), which are also recorded from Antarctica (Shen, 1994; gested by Astrop and Hegna, 2015). Both families are recorded in Africa Everman, 2007; Everman and Stigall, 2007), and the Early Cretaceous with 11 species each. Glyptoasmussia is an interesting genus with five species Nigerestheria lamberti from Niger (Zhang et al., 1976; Shen, species ranging from the Early and Late Triassic to the Late Jurassic in 2003). Furthermore, Roberts et al. (2015) reported the genus Pseu- Africa and with four species from the Lower Jurassic of Antarctica—one destherites and Stigall and Hartmann (2008) the genus Etmosestheria of them (G. luekiensis) in common with Africa (Tasch, 1987). Also, this within the Family Anthronestheriidae. Additionaly, two records of the genus was recorded in the Upper Permian of Australia, Devonian to genus Pseudestherites are mentioned in the Lower Cretaceous from South Upper Triassic in Europe and Russia (Tasch, 1987; Novozhilov, 1958). America (Prámparo et al., 2005; Gallego and Shen, 2006). The problematic genus Pseudestheria (see Cuvelier et al., 2015) is also The Superfamily Estheriteoidea includes mainly a large group of recorded in Africa with 3 species, two of this of late Permian age. This “conchostracans” with radial lirae ornamentation containing all of genus has great diagnostic problems and includes forms with or without known variations from simple lirae to complex chainworks (Zhang ornamentation (see Cuvelier et al., 2015, analyzed four European and et al., 1976; Astrop and Hegna, 2015). In Africa, the Family Asmus- five Russian late Permian species). Pseudestheria is widespread both siidae contains three species in genus Asmussia. These species are stratigraphically and paleogeographically, recently is re-studied and known from Upper Triassic to Lower Cretaceous (Tasch, 1987). This evaluated as an important Paleozoic biostratigraphic marker (Schneider genus is also known from Lower Jurassic of Antarctica and Upper and Scholze, 2018; Scholze et al., 2018). Pseudestheria lepersonnei is the Jurassic of South America (Tasch, 1987). The Family Fushunograptidae only species of this group recorded in two geographic locations—D.R. includes many of the species previously included in the Family Lioes- Congo (Late Triassic) and Angola (Late Cretaceous)—but the older age theriidae (sensu Tasch, 1969 not in the sense of Kozur et al., 1981), and and both stratigraphic sources are still dubious. The genus Orthothemos now corresponds to diagnosis presented by Astrop and Hegna (2015, recorded in the Upper Triassic of Africa is also mentioned from the carapace oval, elliptic or subcircular (cyziciform to telliniform); orna- Upper Permian (3 spp.) and Upper Triassic of South America (2 spp.) mented with fine radial ridges in the growth bands; radial ridges simple and Upper Permian and Upper Cretaceous of Russia (3 spp. belonging to or sometimes branched into a dendritic sculpture, but not reticulated). genera Palaeorthothemos and Sphaerorthothemos). As mentioned above, In the present analysis, we identify nine species as belonging to an other related forms belong to Aquilonoglyptidae, and Aquilonoglypta is unknown genus — six of them were previously assigned to “Lioestheria” recorded in the Lower Triassic of Africa and Upper Permian, and and three others to “Bairdestheria”(Tasch, 1987). The latter genus is a Triassic of Europe and China. genus name with a largely ignored long history, one that probably Some eosestherioid taxa (Eosestheriidae and Anthronestheriidae) needs a thorough reevaluation (Tasch, 1969). These nine species have a have been recorded in the Southern Hemisphere from the Upper long stratigraphic range, from the Early Devonian to the Late Cretac- Triassic of South America (Gallego, 2010), and Lower Jurassic of Ant- eous. Another important component of the superfamily is the Orthes- arctica (Shen, 1994), originally reported by Zhang et al. (1976) from theria-Migransia complex (Shen, 2003). Three species of Orthestheria-

9 O.F. Gallego, et al. Journal of African Earth Sciences 161 (2020) 103648

Migransia from the Upper Jurassic and Lower Cretaceous in Africa are Palaeolimnadia, and Leaia. Besides to each of the four southern con- known (Shen et al., 2004). Also, around forty-five probable species of tinents: Pseudoasmussiata (all except Australia) and Palaeolimnadiopsis estheriteoids group are mentioned from the Permian and Upper Triassic and related genera (all except Antarctica) and to each of three and two (few) and Upper Jurassic to Lower Cretaceous of South America, Aus- continents: Gabonestheria (South America, Africa, India) as well as tralia, India and Antarctica (Tasch, 1987). Estheriella and related genera (Africa, India). The Suborder Estheriellina and the Family Estheriellidae have in- Cyclestherida. Until now, according to Schwentner et al. (2013), teresting records from Africa. Around six species of the genus Estheriella extant Cyclestherida is thought to be represented by a single species and one endemic genus Nyasaestheriella were reported from the Upper only, Cyclestheria hislopi (Baird, 1859), which occurs circumtropically Carboniferous (1 sp.) to Upper Jurassic (2 spp.) with four Triassic (Olesen et al., 1996). Schwentner et al. (2013) however, mentioned that species. The first genus occurs frequently in Europe and North America, Cyclestherida contrast starkly with its putative sister taxon, cladocerans and also in few localities of Gondwana, solely in Africa and India. The (Daphnia, etc.), in terms of diversity and morphological disparity and estheriellid Lioleaiina and also Estheriella are common during the Early proposed that Cyclestheria has three cryptic species in the Southern Triassic in the northern hemisphere (Kozur and Weems, 2010); how- Hemisphere based on molecular studies. In this sense, the assignment of ever, the species recorded from Gondwana are not shared with any fossil forms to the Suborder Cyclestherida is still doubtful, since, as in European or North American taxon. Ghosh (2011) described several all cases of fossil “conchostracans”, the only preserved remains are their estheriellids species from India with endemic forms like Cornutesther- carapaces, lacking these in their majority soft remains. Therefore, fossil iella. Regarding the records of the other species of the Suborder Es- forms assigned to the genus Cyclestherioides are assigned to the Sub- theriellina, the distribution of eight afrograptid species (Liao et al., order Cyclestherida based exclusively on the carapace morphology. 2017) was reported from the Upper Jurassic to the Lower Cretaceous in Fossil Cyclestherida assigned to the extinct taxon Cyclestherioides are Africa, South America and Europe. Afrograptidae comprises five genera known from the Upper Permian in Australia (Raymond, 1946) and (13 spp.) and two of them (Afrograpta, and Camerunograpta, with 2 spp.) Chile (Gallego and Breitkreuz, 1994), Lower Triassic of India (2 spp., are found only in Africa, Graptoestheriella (3 spp.) only found in South Tasch, 1987), Lower Jurassic of Antarctica (Tasch, 1987), the Cretac- America, Surreyestheria (1 sp.) and Lahuerguinagrapta (1 sp.) only found eous in Japan (sensu Chen, 1996) and the mid Eocene in Wyoming, USA in Europe and Congestheriella (6 sp.) was found on three continents (Shen et al., 2006), suggesting that this taxon had a Pangean distribu- (Table 2, Supplementary data). tion as proposed by Schwentner et al. (2013). Nevertheless, Kobayashi The Suborder Leaiina is widespread in Paleozoic deposits from (1954) mentioned a Lower Devonian to Middle Cretaceous distribution Europe, Asia, America, Antarctica, India and Australia. The Family for the genus Cyclestherioides, with around " … 15 species from various Leaiidae occurs widespread in the Middle Devonian to the Upper places and ages, two-thirds of which are in the northern continents and the Permian deposits and they are recorded in North and South America, rest in the southern continents; one-third of the species reported from the India, Australia, Antarctica, China, Siberia and also Central Asia Palaeozoic and the remainder from the Mesozoic era. Eifelian grossi is the (Kazakhstan) and Europe (Raymond, 1946; Novozhilov, 1952, 1956; oldest, and none is known from the later Devonian or from the early Car- Zhang et al., 1976; Chen and Shen, 1985; Ghosh et al., 1987; Jones and boniferous”. Novozhilov, 1958b assigned one Permian species from Chen, 2000; Ferreira-Oliveira and Rohn, 2010). In Africa, this group is Russia to the extant genus Cyclestheria. The low fossil record of Cy- represented by the families Leaiidae (Leaia 1 sp. and Hemicycloleaia 8 clestherida with only five species ranging from the Permian to Jurassic spp.) and Rostroleaiidae (Rostroleaia 1 spp.) and is widespread, parti- from the Southern Hemisphere is remarkable by the more diverse cularly, in Carboniferous and Upper Permian sequences where the Northern Hemisphere records as suggested by Kobayashi (1954) and greatest number of leaiid species (10 spp.) are found (Tasch, 1987). the low Cenozoic to Recent diversity around the Equatorial distribution Southern Hemisphere leaiid faunas show clear differences between the of these taxa (Schwentner et al., 2013). No Cyclestheria specimens from record of South America (only 5 Permian spp., Hemicycloleaia 2 spp., Africa were available for the study of Schwentner et al. (2013), but and Monoleaia 1 sp. and the doubtful genera Acantholeaia 1 sp. and given the results, that continent too may be home to yet another Cy- Unicarinatus 1 sp., these last two forms maybe pelecypods; Rohn, 1989) clestheria species. The results of Schwentner et al. (2013) suggest that a and Antarctica (Leaia 1 sp.). On the other hand, the great diversified combination of vicariance due to the break-up of Gondwana and later faunas from Australia (Hemicycloleaia 7 spp., Rostroleaia 1 spp. ac- intercontinental dispersal between Asia and Australia was responsible cording to the synonymy of Jones and Chen, 2000; or sensu Tasch, 1987 for shaping the current circumtropic distribution of Cyclestheria species, Hemicycloleaia 23 spp., Leaia 1 sp.) and India (Leaia 3 spp., Hemi- dating the age of crown group Cyclestherida to the Cretaceous. Also, cycloleaia 6 spp., Monoleaia 1 sp., Rostroleaia 4 spp. sensu Ghosh, 2011). those authors mentioned that depending on the phylogenetic position This information suggests close biogeographic relationships between and overall genetic differentiation of African populations, the age ofthe Africa, India and Australia and suggest there should be common species crown group of Cyclestherida may actually be even greater. This would among them, assuming that leaiids used dispersal mechanisms similar be consistent with the Northern Hemisphere Devonian to Carboniferous to modern clam shrimps. records of the Cyclestherida stem group plus species assigned the extant genus Cyclestheria (Novozhilov, 1958b). 5. Discussion According to Tasch (1987), the dispersal of Cyclestherioides between Africa and Antarctica appears to be one possibility in light of their 5.1. Paleobiogeographic implications Jurassic records, or both the African and Antarctic occurrences of Cy- clestherioides species may have originally derived from the Early Triassic In this section, we discuss potential dispersal routes of “con- population of India. Also, previous records from the Permian of Chile chostracans” between African and other paleocontinents informed by and the Lower Paleozoic of Europe support, as suggested by Schwentner taxomic and paleogeographic relationships described above. We re- et al. (2013), a Pangean distribution and its dispersal from South cognize the incompleteness of “conchostracans” fossil record, and pre- America and Europe before Permian times. sent this discussion as a qualitative hypothesis to help inform future, Whatever dispersal tracks prevailed, the spread of Cyclestherioides- potentially more quantitative analyses. The information was treated Cyclestheria bioprogram to South America, India, Africa, Australia, and according to systematic groups. Antarctica actually occurred (Schwentner et al., 2013). It is evident that Tasch (1987, now actualized with our database) established that his dispersal of species eggs of the five genera whose fossil distribution is so called “bioprograms” of five “conchostracan” genera were dispersed reviewed above, occurred to each of the five continents during the to all five continents of the Southern Hemisphere during some portions Permian-Triassic and Jurassic in the Southern Hemisphere. Multiple of Paleozoic/Mesozoic times as, Cyclestherioides, Cornia, Estheriina, examples of such distributions of other “conchostracan” genera to four

10 O.F. Gallego, et al. Journal of African Earth Sciences 161 (2020) 103648 or three of the southern continents are reviewed below. The Family Lioestheriidae (sensu Kozur et al., 1981) is well-re- Spinicaudata. Within this group, the Vertexioidea-Palaeolimnadiidae presented in Africa by the genera Echinestheria, Cornia (Fig. 3C), Ga- group were widespread records during the late Permian-Triassic, although bonestheria, Falsisca and Palaeestheria ranging from the Carboniferous to subsequently they became restricted to the Jurassic-Cretaceous of the the Early Cretaceous, suggested that this continent played an important Southern Hemisphere. Two genera, Palaeolimnadia and Estheriina, dominate role in the diversification and evolution of this spinicaudatan group. the fossil records of the group. The late Permian (Australia, India, and South These taxa, specially Gabonestheria, Falsisca and Palaeestheria require America) Palaeolimnadia and Estheriina species are the oldest, and when revisions of their type material (because Gabonestheria might be sy- added to the record of Triassic forms; Australia and Africa share the most nonym to Cornia and Falsisca might be synonym to Palaeolimnadiopsis, diverse fauna with twenty-one species (four species in common). This si- which belongs to Palaeolimnadiopseidae instead of Lioestheriidae). The tuation prevents us from discovering in what sense and from when and shared records between South America, India and Australia (Permian- where the palaeolimnadiids were distributed in the southern region of our Triassic Echinestheria, Cornia and Gabonestheria species) and Early Jur- planet. Only the record of three Permian species in Australia and also in assic species from Antarctica, suggest a long evolutionary history in the South America suggests a probable origin of the group in the southern southern continents. This ancient past is also, supported by the record terranes, but less is known on their history in the northern hemisphere. of Early Triassic Indian endemic forms, such as Turricornia and In- Afrolimnadia is recorded in both the Lower Triassic of India and the Upper domonocarina. Also, Vertexia, Protomonocarina and Curvacornutus share Triassic of Africa suggesting other close affinities to be found in Gondwana. records in the Upper Paleozoic and Triassic from North America and The Early Jurassic and Early Cretaceous species distributed in Antarctica Europe, which shows close biogeographical relationships. (11 sp.) and South America (7 spp.), must have used Africa as an important The Family Ulugkemiidae is another example of probable Paleozoic bridge between the other southern continents. taxon (Middle Devonian - late Permian, Russia) that migrated from Tasch (1987) mentioned several points about the spread of the es- northern to southern region through the Africa continent with members theriinid bioprogram (Estheriininae) that merit further comment, in the Lower Triassic of Africa and Middle-Upper Triassic of South herein augmented with new data: 1. South America and India (Ghosh, America (Gallego and Melchor, 2000). Those authors proposed two 2011) are the only two southern continents having late Permian es- possible ways to the dispersal of this group as the transport on the theriinids, the geologically oldest known representatives of the genus. tetrapod bodies with aquatic habits and the trans-equatorial winds as a 2. Accordingly, dispersal of estheriinid eggs from the Upper Permian to result of the monsoonal climate of the Triassic Pangea. the Lower Triassic seems possible in both India (Ghosh, 2011) and Other records that support the north-south exchange correspond to Brazil areas, although diachronous. As noted above, the Canning Basin the species level analysis with Cornia? limbata (Lioestheriidae, Fig. 3D) of Western Australia and the Bowen Basin of eastern Australia also had and ?Euestheria simoni (Fig. 3E, see Hennion et al., 2015), both Penn- Early Triassic estheriinids. If it is quite speculative, Tasch (1987, p. 140) sylvanian European species recorded also in the Carboniferous of said: “Because the Blina Shale of the Canning Basin is geologically younger Africa. Schneider and Scholze (2018) mentioned the first one as Pseu- than the base of the Rewan Formation of the Bowen Basin, dispersal would destheria limbata that belong to the Pseudestheria limbata–Pseudestheria have been from east to west” (see Tasch in Tasch and Jones, 1979c, p. rimosa–Lioestheria form Köllerbach assemblage zone from the Stepha- 37–38). That leaves the question of how the estheriinids reached the nian A and ?Stephanian B. ?Euestheria simoni constitutes an excellent Bowen Basin from India or Brazil. 3. The Upper Triassic of Angola with fossil guide of the Westphalian C-D Anomalonema reumauxi-Pseu- the record of the Early Triassic Australian species (Estheriina (Nudusia) destheria simoni Assemblage-Zone (Schneider et al., 2005, in Cuvelier cf. rewanensis) could have received dispersed estheriinids from Aus- et al., 2015). Scholze et al. (2018) analyzed the value of the genus tralia, but an exchange with South America or India (Ghosh, 2011) Pseudestheria but did not comment on the new combination of P. limbata cannot be ruled out. 4. The Upper Triassic of Africa (Angola) is a pos- (among other species) cited in Schneider and Scholze (2018), which we sible original source of the Antarctic's Lower Jurassic estheriinids, but a believe is not valid as a formal nomenclatural act. relationship with South America, India or Australia should be checked. The Family Euestheriidae is one of the best represented records in On the other hand, data on a Lower Jurassic Antarctic links is presently Africa. There are nineteen (19) species from the Upper Carboniferous to lacking. 5. India, with its several Lower Jurassic estheriinid species, is a Upper Cretaceous, with nine (9) Triassic species from South American more viable option than Angola for the source of Antarctic estheriinids (Gallego, 2001a, b). Also the family is represented by the questionable (see Tasch, 1987, Frontispiece, pag. ii, Gondwana Fossil Conchostracan genus Pseudoasmussia? (see Cuvelier et al., 2015; Hennion et al., 2015) Distribution). It is worth nothing that India and Antarctica, respec- with three (3) species ranging from the Late Jurassic to the Early Cre- tively, have the only Early Jurassic estheriinids in the reconstructed taceous. The occurence of Euestheria minuta, E. mangaliensis and E. continental assemblage. 6. Dispersal of Antarctic Jurassic estheriinids forbesi in the Upper Triassic of Africa and the Middle - Upper Triassic in presumably was the ultimate source of Brazilian forms of Early Cre- South America (Gallego, 1992, 2001a, b) supports a close relationship taceous age, although the time hiatus still needs to be bridged. Never- between both continents (Martins-Neto et al., 2003). Euestheria man- theless, South America has a great record of this group from the Upper galiensis is another interesting species that shows a south-north trend Permian, Upper Triassic and Upper Jurassic that probably gave rise to with its original record from the Lower Triassic of India but with the the Cretaceous forms. oldest record in the Upper Permian (data came from Bond, 1955 and The Vertexioidea-Palaeolimnadiopseidae is a very interesting group Tasch, 1987, p. 57–58, for more information see Supplementary Data) from both a systematic point of view (Astrop and Hegna, 2015) and a Figure and Lower Triassic of Africa. It is a Gondwanan species that dispersed to Multimedia component 4 paleobiogeographical perspective. The most im- South and North America and also to Europe and by other route to portant record of the group is the presence of Pteriograpta reali in Africa and Australia. Nevertheless, as in other “conchostracan” species this one Pteriograpta cf. reali in South American Jurassic-Cretaceous sequences, needs a complete study, focused in it many populations, and also it supporting an Africa-South American bioprovince during this time (Gallego assignment to the genus Magniestheria (see Kozur and Seidel, 1983; et al., 2010). Nevertheless, the Devonian, Carboniferous and Permian re- Scholze et al., 2016). Euestheria minuta is another species that shows a cords in the Northern Hemisphere (Palaeolimnadiopsis carpenteri, P. eifelensis, similar dispersal pattern but mainly for the Triassic times, it has the and others see Raymond, 1946) suggest a northern origin of the group, and oldest record from the Early Triassic of Africa and Middle Triassic of then dispersed to South America and India-Australia during the early-late Europe (sic. Defretin, 1950) and mainly Late Triassic records from Permian. Also, the presence of the endemic species Stanleyviella recorded in South America, Australia, east and southeast Asia, China, Russia, and the Mesozoic of Africa and the absence of Macrolimnadiopsis forms (re- Europe, suggested an East continental-island corridor species (Ec-ic, corded in South America and India) suggest a fair bit of heterogeneity proposed here, Fig. 4). Also, E. minuta dispersed during the Late Triassic among clam shrimp faunas. by the other way via West continental corridor (Wcc, proposed here,

11 O.F. Gallego, et al. Journal of African Earth Sciences 161 (2020) 103648

Fig. 4. Pangean reconstruction (modified from Blakey, 2005) showing schematic diagram of zonal circulation take of Parrish (1990) and Roscher et al. (2011). Arrows are surface winds vectorized according to the reigning general direction based on Roscher et al. (2011).

Fig. 4) reported by records from South America (also with Middle indicates that these taxa spread across Gondwana during Triassic. Triassic records), North America, Greenland and Europe, and then Several African occurrences of species of the Superfamily reaching Antarctica during the Early Jurassic. Other two Euestheria Eosestherioidea provide geographic context for the evolution of the species, E. triassibrevis (Australia) and E. truempyi (Europe, also know as “conchostracan” fauna. The Family Eosestheriidae defined by Zhang Magniestheria Kozur and Seidel, 1983) are shared with the Lower et al. (1976) exhibits high diversity and a broad geographical dis- Triassic record from Africa and suggest a Triassic link across north and tribution but is restricted temporally as mentioned by Astrop and Hegna south areas. Also the species “Estheria” destombesi (Fig. 3F) is recorded (2015). The Late Triassic and the Early Jurassic records of eosestheriids in the Middle-Upper Triassic from Europe and the Late Triassic of extended back the stratigraphical distribution in relation to the original Africa. These exchanges probably continues during the late Mesozoic range of Late Jurassic to Early Cretaceous records from China (Zhang suggested by the record of Jurassic (Mongolia and Transbaikalia) and et al., 1976; Chen and Shen, 1985). Also the African records of this Cretaceous (Africa) species, as Euestheria dahurica. The questionable family from the Upper Triassic, Lower Jurassic and Lower Cretaceous genus Pseudoasmussia (or Pseudoasmusiata Tasch, 1987; see comments (Zhang et al., 1976; Shen, 2003; Everman, 2007; Stigall et al., 2014) in Hennion et al., 2015) also reflects a north-south trend suggested by supported the oldest findings in the Southern Hemisphere mentioned by the record of P. grassmücki from the Triassic of Greenland and three (3) Shen (1994) and Gallego (2010). All these records contribute to suggest species of that genus reported from the Upper Jurassic – Lower Cre- a probable southern origin of the group and the important role of taceous from Africa. African forms to allow these interpretations (sensu Stigall et al., 2014). The Family Orthothemosiidae joined with Aquilonoglyptidae is Furthermore, the recent documentation from the Upper Cretaceous of dubious in validity, but with an important record of eleven (11) species Africa of the genera Pseudestherites by Roberts et al. (2015) and Etmo- in African. Astrop and Hegna (2015) considered both as part of the sestheria (Stigall and Hartmann, 2008) within the Family Anthro- Family Euestheriidae. Glyptoasmussia has it oldest record from the De- nestheriidae bring more evidence about the role of the Gondwanan vonian of Europe and Russia, and according to this finding then spread faunas into the evolution of this group. Through the new records of the to the Upper Permian and Upper Triassic of the northern hemisphere Family Anthronestheriidae from Africa (adding the Early Cretaceous and dispersed to the Upper Permian of Australia (Tasch, 1987). Later, from South America, Prámparo et al., 2005; Gallego and Shen, 2006) this genus was dispersed and diversified with five (5) species that Pseudestherites has a short stratigraphical distribution restricted to the ranges from the Early and Upper Triassic to the Late Jurassic in Africa, Berriasian to Barremian stages from China, now probably extended to and then migrated to the Lower Jurassic of Antarctica where are re- the Late Cretaceous (Africa). corded four (4) species of that genus and Glyptoasmussia luekiensis ori- The Superfamily Estheriteoidea includes mainly a large group of ginally described from Africa. On the other hand, the genus Pseu- “conchostracans” with striated ornamentation with all of known var- destheria (for the meaning of this taxa see Cuvelier et al., 2015 and iations from simple striae to complex chainworks. In this group we Scholze et al., 2018) with two species of late Permian age is also re- included Asmussiidae with three species from the Upper Triassic to the corded in Africa. This genus has a wide spread stratigraphically and Lower Cretaceous of Africa, one species from the Lower Jurassic of paleogeographically, but the meaning of this record needs more studies Antarctica and other one from the Upper Jurassic of South America. As to define its real taxonomic validity (see Scholze et al., 2018). Genera mentioned by Tasch (1987) “… The significant point is that the above like Orthothemos and Aquilonoglypta (Aquilonoglyptidae) are recorded spread of the founder genus, Asmussia Pacht, originally described from the in the Upper Permian and Triassic of Russia, Europe, China and South Lower Devonian of "Livonia" (sic, actually part of Latvia and Estonia) and America and the Late Cretaceous of Russia. Nevertheless, they are only some related genera that evolved from its species, constitutes clear evidence recorded in the Lower and Upper Triassic of Africa, which probably of Mesozoic Southern Hemisphere dispersal to each of the four continents

12 O.F. Gallego, et al. Journal of African Earth Sciences 161 (2020) 103648

(other Lower Devonian forms of Asmussia, as well as Upper Permian species, India and Africa. This group exhibits widespread distributions, parti- are known from the Russian Arctic; see Novozhilov, 1958b)”. According cularly, in Permian sequences where the greatest number of leaiid to this mention, the Asmussiidae is a northern group that reached the species (13 spp.) occur in the Upper Permian of Australia (Tasch, 1987) southern domain during the Pangea times. and India (14 spp., Ghosh, 2011). Sharing with Africa the presence of The African record of the Superfamily Estheriteoidea (and probably genera Leaia, Hemicycloleaia and Rostroleaia. Jones and Chen (2000) Family Fushunograptidae among others) is composed by nine (9) spe- proposed the distribution of Hemicycloleaia mitchelli Etheridge, 1892 cies with six (6) “Lioestheria” and three (3) “Bairdestheria” that range and other species as bipolar distribution of cool waters in the middle- from the Early Devonian to the Late Cretaceous (Tasch, 1987). The late Permian and these “conchostracans” were conspecific in the Jurassic-Cretaceous is the most diverse with five (5) records and the Northern and Southern Hemispheres and dispersed between Siberia and next is the Late Triassic with three (3) records. B. kitariensis and L. le- eastern Australia via China (Jones and Chen, 2000). However, the sothoensis are recorded in two distinct areas Angola-Congo and Lesotho- leaiids found in the Southern and Northern hemispheres did not ne- South Africa respectively. The species L. cassambensis and B. mawsoni cessarily have a dispersal route through the China-Australian pathway have a real paleogeographical meaning, because both are recorded in (East continental-islands corridor – Ec-ic, defined here, Fig. 4); for ex- the Late Jurassic and Early Cretaceous from Africa and South America ample, the north African region is another possibility (Ferreira-Oliveira supporting the ASA province (Gallego et al., 2010). The five (5) Late and Rohn, 2010). Africa leaiid records have three (3) and four (4) Jurassic-Late Cretaceous species of Orthestheria-Migransia and related species of Hemicycloleaia from Carboniferous and Permian respectively, forms from Africa show close relationships with the around fourty (40) supporting as possible distribution of this genus through Africa to the species of estheriteoids group of the approximately same age from the rest of the southern hemisphere (West continental corridor – Wcc, de- rest of southern continents. The presences of the Asiatic genera (Or- fined here), which was high worldwide during most of the late Paleo- dosestheria and Cratostracus) from the Early Cretaceous of northern zoic (Fig. 4). This kind of proposal is in concordance with the revision Africa also suggest close affinities with this fauna. Furthermore sup- and suggested ideas given by Jones and Chen (2000) for some index porting previous ideas about the presence of a series of lacustrine sys- leaiids species Hemicycloleaia andersonae (Tasch, 1979), H. grantrangicus tems interconnected the north of South America with Southeast Asia (Tasch, 1979) and Rostroleaia carboniferae (Tasch, 1979); and the based on selected groups (Musacchio, 2001; Martins-Neto, 2002, 2006; known late Permian taxa to three species as, Hemicycloleaia mitchelli Martín-Closas and Wang, 2008; Buscalioni and Poyato-Ariza, 2016, (Etheridge, 1892), H. discoidea (Mitchell, 1925), and H. deflectomarginis Fig. 8; Figure Multimedia component 4). (Tasch, 1979; see Tasch and Jones, 1979). Estheriellina. Comprise two interesting groups from the biostrati- graphical and paleobiogeographical point of view: the superfamilies 5.2. Biogeographic links through time Estheriellioidea and Afrograptioidea. The Estheriellidae has a wide- spread record during late Paleozoic and Triassic representing an im- The close relationships between Africa and the other four portant stratigraphic tool. The genus Estheriella occurs frequently in Gondwana components along the geological time were clearly defined Europe, and also in few localities of only North America by pioneers like Alexander Du Toit and Alfred Wegener in the first (Acadiestheriella), Africa and India. Six African species of the genus are decades of nineteenth century. Wegener (1912) used the record of fossil known that clearly differs from the Early Triassic estheriellid fauna emblematic taxa (as Lystrosaurus, Mesosaurus, Glossopteris, etc.) to de- from India composed by three (3) endemic species and five (5) ones of monstrate the paleogeographic connection between southern con- Estheriella. Also differs from the northern estheriellid faunas that com- tinents. During the Carboniferous, the supercontinent Pangea was prises the estheriellid Lioleaiina, Acadiestheriella and the index species. completed, and it has often been suggested that were few barriers to E. costata and E. nodosocostata are common during the Early Triassic of dispersal, producing a strongly cosmopolitan fauna at least the family the northern hemisphere (Kozur and Weems, 2010), however, the levels (i.e., Colbert, 1973; Olson, 1996). species recorded from Gondwana are not shared with Europe or North America. Kozur and Weems (2010) mentioned that E. costata and E. 5.2.1. Biogeographical controls nodosocostata have a regional distribution ranging from the Germanic For all this information mentioned above, the supercontinent Basin, across the Russian Platform and Gondwanan India to Angola. Gondwana was a place that fostered dispersal of continental organsims Ghosh (2011) described eight (8) Triassic estheriellids species from like "conchostracans". The paleogeography of Gondwana was similar India, with endemic forms like Cornutestheriella and two (2) new sub- from the Late Permian through the Middle Jurassic. Through much of genus and four (4) new species of Estheriella. Regarding Afrograptidae its history, Gondwana probably was large enough to create the strongly records, the distribution of their species were reported from the Jurassic seasonal circulation pattern known as a monsoon. The large size of the and the Cretaceous in Africa, Europe and South America. Afrograptidae continent also would have meant that the interior of the continent was comprise five (5) genera and three (3) of them are found in Africa, two arid. Even in the absence of mountains, moisture-laden (i.e., warm) (2) in South America and two (2) in Europe (Table 2). This record winds flowing landward lose their moisture over land because heating supports the idea of West continental corridor (Wcc, defined here) as drives the air masses upward (in winter, the moisture in landward- important migration route where Afrograptidae distributed both in the flowing air does not condense unless the air is forced upwardby Northern and Southern Hemispheres and the establishment of the ASA mountains or by a warm air mass from another source). Thus, in low biogeographic province in the Jurassic to Early Cretaceous (Shen, 2003; latitudes, rainfall not only would have been seasonal, but also confined Gallego et al., 2010; Gallego and Martins Neto, 2006). Finally, Esther- to coastal regions (Parrish, 1990). Wind direction vectors proposed by iellidae show a widespread across the Northern Hemisphere but is re- Roscher et al. (2011) shown a predominance of winds from high to low stricted for the Gondwana supercontinent with only two records from latitudes. The higher annual rainfall in western most tropical Pangea is India and Africa. Conversely, Afrograptidae has a great record in the caused by westerly equatorial winds hitting the continent from the Southern Hemisphere and up to now a low record in Europe (England, Panthalassan side. The two mid-latitude precipitation regions are Bulgary, and Spain; Gallego et al., 2018) marked a different way of bound to the westerly wind belt at about 50° north and south respec- faunal exchange between northern and southern domains. Other rea- tively. The intensity within these belts decreases from the west-coasts sons for this low record of Afrograptidae in Europe can be a con- towards the continental interiors of Siberia, India and Australia sequence of predominance of marine facies (e.g., mainly marine de- (Roscher et al., 2011). posits). On the other hand, dispersal barriers developed from increased Leaiina. As previously mentioned, this group is widespread in mountain-bulding (e.g. Appalachian orogeny) and due to environ- Paleozoic deposits from Europe, Asia, America, Antarctica, Australia, mental changes probably played an important role in the distribution of

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“conchostracans”. These physical and environmental barriers played a The presence of afrograptid “conchostracans”, as well as other important role in the distribution species of plants (Cleal et al., 2009) groups, shows close paleogeographic relationships between northern and vertebrates (Brocklehurst et al., 2018). Recently, provincialism and South America and Central Africa during the Late Jurassic to Early faunal variation according to palaeolatitude has been identified in Cretaceous (Carvalho, 1993; Rohn and Cavalheiro, 1996; Arai and several vertebrates (Modesto and Rybczynski, 2000; Sidor et al., 2005; Carvalho, 2001; Cunha Lana and Carvalho, 2002; Rohn et al., 2005). Ezcurra, 2010). Shen (2003) (in Gallego and Martins-Neto, 2006) proposed that during In this way, the wind patterns, physical and climate barries, prob- the Late Jurassic and Early Cretaceous existed a biogeographic pro- ably, generated also a division of "conchostracan" fauna. The fossil re- vince, including Africa-South America (ASA) represented by Afro- cords of the Estheriellina group support this idea, where Estheriella graptidae fauna (Afrograpta, Camerunograpta, Graptoestheriella and occurs frequently in Europe and North America and the afrograptids Congestheriella) adding the joint record of Pteriograpta cf. reali, “Lioes- show a great record in the Southern Hemisphere. However, the wind theria” mawsoni and “Lioestheria” cassambensis in both areas, Brazil and patterns from Africa- North of South America (mid-latitude) to central Africa. Moreover, new evidence provided by the record of Antarctic-Australia (high latitude) generated dispersal "conchostracan" Congestheriella (Gallego et al., 2010) combined with the record of Or- fauna into the Gondwana, as case of the estheriinids and palaeolima- thestheria (Migransia) ferrandoi (a "Bairdestheria" mawsoni - "Lioestheria" diids. Although, some species of Leaiina exhibit widespread in Pangea cassambensis related form) tentatively extends this paleobiogeo- or bipolar distribution as Hemicycloleaia mitchelli where the dispersal graphical province to northwestern (Venezuela) and southern (Argen- route through the China-Australian vias (Jones and Chen, 2000) or tina and Uruguay) South America during the Middle to Late Jurassic. through Africa (Ferreira-Oliveira and Rohn, 2010) could had possible. Recently, the record of afrograptids like Camerunograpta-Graptoesther- In addition, the record of the dispersal of the "conchostracan" species iella species and a new form in the Lower Cretaceous of Europe (Liao was different across geological time. et al., 2017; Gallego et al., 2017, 2018) spread the geographical dis- tribution and the influence of the African fauna to the Northern 5.2.2. Late Paleozoic-Triassic Hemisphere. Other Jurassic-Cretaceous records of fushunograptids, The African “conchostracan” records suggest four different faunal anthronestheriids and eosestheriids suggest another connection ex- interchanges (Tables 1 and 2; Supplementary data) between Africa and change with Europe, Asia, Antarctica and southern South America each one area, Australia–Europe–South America and India marked by (Stigall and Hartmann, 2008; Stigall et al., 2014). Loxomegaglyptid different emblematic taxa. records in the Upper Cretaceous in Africa and its relation with the The Australian–Africa fossil invertebrates interchanges were con- Triassic and Lower Cretaceous records from South America would be firmed by the presence of leaiids during the Carboniferous–Permian of explored in the future and also suggest a close interchanges between Africa, not compared with other southern records. During the late both areas in one direction only (from South America to Africa). Pa- Permian and Late Triassic the presence of palaeolimnadiids in both laeolimnadiids and palaeolimnadiopseids also show a low diversity in Africa and Australia (sharing, Palaeolimnadia banksi, P. cf. wiana- Africa but an interesting relation with the Early Jurassic faunas of India mattensis, P. cf. glabra, Estheriina cf. rewanensis) support that largest and Antarctica and the Early Cretaceous fauna from South America. exchange between both continents and previous to the post Middle Jurassic spread of Africa through India and Antarctica. Tasch (1987) 6. Conclusions mentioned that the Triassic of Angola could have received dispersed estheriinids from South America. The great records of euestheriids and The new comprehensive compilation of “conchostracan” ocurrences the presence of the lioestheriid Gabonestheria gabonensis characterize increase our knowledge on the diversity and abundance of the fossil this time and suggest a close African-South America and Europe ex- record of “conchostracans” from Africa and even from the other change. On the other hand, Euestheria simoni (or Pseudestheria simoni Gondwana continents. In particular, related to the scope of this work according to Schneider et al., 2005) is an index taxa for the Carboni- we confirm previous ideas on the role of Africa in the diversification ferous of Europe and of course with Africa that shows close relation- and dispersal of the “conchostracan” faunas to and from the African ships between both continents that conforms the Euroamerican bio- continent. graphic province (also supported by insect fauna see Belahmira et al., The history of “conchostracan” fossil groups in Africa and 2019). Furthermore, Cornia limbata (or Pseudestheria limbata sensu Gondwana can be characterized into two main phases. 1) the Devonian Schneider and Scholze, 2018) recorded also in the Carboniferous of to Triassic as a lapse of time with a strong composition of leaiids, Africa is another species with the same significance as E. simoni. lioestheriids, palaeolimnadiids and euestheriids. The African “con- However, these are taxa of doubtful validity the genus Falsisca and the chostracan” records suggest four different faunal interchanges between ulugkmiids are other taxa that show this affinity. The record of Es- Africa and each one area, Australia - Europe - South America and India theriellidae from the Upper Carboniferous to Upper Permian and Lower marked by different emblematic taxa. In this time span also, the other to Upper Triassic from Africa suggest a close relation with the Early important groups for paleobiogeographic meanings for Pangea are in- Triassic fauna of India. Kozur and Weems (2010) mentioned “… that sects and bivalved mollusks. 2) The Jurassic to Cretaceous times as a this very wide distribution of many species of conchostracans in both in the time with a great number of afrograptids, fushunograptids, palaeo- northern hemisphere and across large parts of Gondwana ended during the limnadiopseids, palaeolimnadiids, loxomegaglyptids, anthronestheriids Carnian. Upper Triassic conchostracans of Argentina, Brazil and Chile are and eosestheriids. The “conchostracan” records suggest five different very different from the conchostracan faunas of the northern hemisphere, faunal interchanges between Africa and each one area, South America - and only in northwestern Africa do the same conchostracans occur as in the Europe - Asia - India and Antarctica marked by different emblematic northern hemisphere..”. Some family-level lineages whose diversity is taxa. centered in the Paleozoic survived the end-Permian event into the This new data compilation can serve as a springboard for future earlier Mesozoic, such as the Family Aquilonoglyptidae and Peri- studies including quantitative analysis. Adequate identification of some limnadiidae. species and data on age of deposits are needed to further understand the paleogeographic relationships of "conchostracans". 5.2.3. Jurassic – Cretaceous The “conchostracan” records suggest five different faunal inter- Acknowledgements changes (Table 2; Supplementary data) between Africa and each one area, South America - Europe - Asia - India and Antarctica marked by This work was supported by the Consejo Nacional de Investigaciones different emblematic taxa. Científicas y Técnicas [CONICET-PIP-11220150100117CO]; Secretaría

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General de Ciencia y Técnica- Universidad Nacional del Nordeste [SEGCyT- Nat. Hist. 15, 1–12. UNNE PI-Q006-2014 and PI-18Q005], Argentina. Also thanks to the Musée Chen, P.J., Shen, Y.B., 1985. An Introduction to Fossil Conchostraca. Science Press, Beijing, pp. 241. Gosselet, Lille (Musée d’Histoire Naturelle – Musée de Géologie) and Lille Chernyshev, B.I., 1930. Estheria from Siberia and the far east. 49 Central Geological University – Sciences and Technologies (France) and National Museum of Prospecting Bureau, Bulletin 85-70. Natural History (USA) for allow us to publish specimens loaned in each Cleal, C., Opluštil, S., Thomas, B., Tenchov, Y., Abbink, O., Bek, J., Zodrow, E., 2009. Late moscovian terrestrial biotas and palaeoenvironments of variscan euramerica. institutions. The authros are thanked to Dr. Joerg Schneider (TU Netherlands Journal of Geosciences - Geologie En Mijnbouw 88 (4), 181–278. Bergakademie Freiberg), Dr. Liao Huanyu (Yunnan University, Yunnan https://doi:10.1017/S0016774600000846. Province, China) and two other reviewers for the improved suggestions to Colbert, E.H., 1973. Continental drift and the distributions of fossil reptiles. Proc. Natl. our manuscript. Acad. Sci. India 1, 395–412. Colin, J.-P., Dépêche, F., 1997. Faunes d’ostracodes lacustres des bassins intra-cratoniques d’age albo-aptien en Africa de l'Ouest (Cameroun, Tchad) et au Brésil: considérations Appendix A. Supplementary data d’ordre paléoécologique et paléobiogéo- graphique. Afr. Geosci. Rev. 4, 431–450. Colin, J.-P., Brunet, M., Congleton, J.D., Dejax, J., Flynn, L.J., Hell, J., Jacobs, L., 1992. Ostracodes lacustres des bassins d’âge crétacé inférieur du Nord du Cameoun: hama- Supplementary data to this article can be found online at https:// Koussou, Koum et Barbouri-Figuil. Rev. Paléobiol. 11, 357–372. doi.org/10.1016/j.jafrearsci.2019.103648. 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