Cretaceous Research 154 (2024) 105752 Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes A bioprovince for the BarremianeAptian charophytes of the Central Tethyan Archipelago Alba Vicente a, b, c, d, Josep Sanjuan b, c, *, Jordi Pe rez-Cano b, c, d, e, Khaled Trabelsi f, g, h, Carles Martín-Closas b, c a Instituto Polit ecnico Nacional, Avenida Instituto Polit ecnico Nacional s/n, Playa Palo de Sta Rita, La Paz, Baja California Sur, 23096, Mexico b Departament de Dina mica de la Terra i de l'Ocea , Facultat de Ci encies de la Terra, Universitat de Barcelona (UB), Martí i Franques s/n, Barcelona, Catalonia, 08028, Spain c Institut de Recerca de la Biodiversitat (IRBio), Universitat de Barcelona (UB), Barcelona, Catalonia, 08028, Spain d Institut Catala de Paleontologia Miquel Crusafont, ICTA-ICP Building, Universitat Auto noma de Barcelona, c/de les Columnes s/n, Cerdanyola del Vall es, Catalonia, 08193, Spain e Departament de Mineralogia, Petrologia i Geologia Aplicada, Facultat de Ci encies de la Terra, Universitat de Barcelona (UB), Martí i Franqu es s/n, Barcelona, Catalonia, 08028, Spain f Department of Earth Sciences, Faculty of Sciences of Sfax, University of Sfax, Sfax, 3038, Tunisia g LR18ES07, Department of Earth Sciences, Faculty of Sciences of Tunis, University of Tunis El Manar II, Tunis, 1068, Tunisia h Department of Geology, Faculty of Earth Sciences, Geography and Astronomy, University of Vienna, Josef-Holaubek-Platz 2, Vienna, 1090, Austria a r t i c l e i n f o a b s t r a c t Article history: A comprehensive bibliographic compilation of dozens of worldwide records of charophytes from the Received 2 May 2023 Barremian to early Aptian interval has been revisited aiming to gather the regional and global Received in revised form distribution data for the clavatoracean family. The compiled data were studied with the palaeontological 17 October 2023 statistics software PAST (PAleontological STatistics), using a cluster analysis. All analyses of the biogeo- Accepted in revised form 18 October 2023 Available online 23 October 2023 graphic data revealed a consistent separation of the Central Tethyan Archipelago (CTA) from other palaeogeographic areas, mainly the neighbouring Eurasian and American landmasses, based on the species richness of Family Clavatoraceae. Keywords: Charophyta The Central Tethyan Archipelago (CTA) is hence, proposed as a distinct bioprovince for Early Creta- Lower Cretaceous ceous charophytes, called the Clavatoracean Bioprovince, based on the abundance and especially the Mesogea biodiversity of clavatoracean taxa (up to 26 taxa). Within the Archipelago, species distribution primarily Tethys shows a latitudinal pattern suggesting that the charophyte floras were subjected to climatic control. Mesozoic However, longitudinal relationships between areas in the eastern or western parts of the CTA were also Island biogeography found. In spite of these internal subdivisions of the Clavatoracean Bioprovince, the flux of populations between the islands must have been sufficiently strong to maintain taxonomic homogeneity within the archipelago throughout the time interval analysed. These biogeographic patterns are compared with those of non-marine ostracods for the same time interval, showing strong similarities and suggesting that they may be equivalent for other non-marine benthic organisms. © 2023 Elsevier Ltd. All rights reserved. 1. Introduction fructifications bear sufficient vegetative and reproductive charac- ters to determine the species with much more confidence than The clavatoraceans (Family Clavatoraceae Pia) are an extinct, other fossil charophytes (e.g. Grambast, 1974; Martín-Closas, mainly Cretaceous group of charophytes that produced complex 2000). Several factors indicate that clavatoraceans represent an calcified fructifications called utricles. These biomineralized excellent group for palaeobiogeographic studies. First, Clavator- aceae utricles are abundant microfossils that are easy to recover from the bearing rocks and to prepare for study. Second, some * Corresponding author. Departament de Dina mica de la Terra i de l'Ocea , Fac- species show a wide palaeogeographic distribution (continental, ncies de la Terra, Universitat de Barcelona (UB), Martí i Franque ultat de Cie s s/n, sub-cosmopolitan, and cosmopolitan) during the Early Cretaceous Barcelona, Catalonia, 08028, Spain. (e.g. Martín-Closas and Wang, 2008; Martín-Closas, 2015), coupled E-mail address: josepsanjuan@ub.edu (J. Sanjuan). https://doi.org/10.1016/j.cretres.2023.105752 0195-6671/© 2023 Elsevier Ltd. All rights reserved. A. Vicente, J. Sanjuan, J. Perez-Cano et al. Cretaceous Research 154 (2024) 105752 with a relatively high evolutionary rate, rendering them very data to help to better define the area where current biozonations useful in biostratigraphy (rock age determination) and strati- can be applied. graphic correlation worldwide (e.g. Musacchio, 1973; Grambast, 1974; Wang and Lu, 1982; Riveline et al., 1996; Li et al., 2016; 2. Materials and methods Perez-Cano et al., 2022a). This application in geology, has yielded large quantities of presence data on many continents. Third, some 2.1. Data acquisition clavatoracean species evolved following anagenetic processes, i.e. gradual evolution of intraspecific lineages (e.g. Grambast, 1974; The dataset used in this study consisted of bibliographic Martín-Closas, 1996; Vicente and Martín-Closas, 2013; Pe rez-Cano references and own data on the occurrence of clavatoracean et al., 2020), providing significant evolutionary information on the charophytes during the Barremianeearly Aptian. This time interval historical biogeography of each species (e.g. Martín-Closas and was selected because of its rich charophyte fossil record worldwide Wang, 2008). Fourth and lastly, during the Early Cretaceous clav- and because it coincides with the maximum species richness of atoraceans thrived in a wide range of non-marine environments clavatoracean charophytes (Martín-Closas and Serra-Kiel, 1991). such as permanent alkaline lakes, temporary floodplain ponds, and The database contains the taxonomy, the age of the deposit, the brackish marshes (e.g. Climent-Dome nech et al., 2009; Vicente and locality, the geographic area or sedimentary basin, the country, and Martín-Closas, 2013; Pe rez-Cano et al., 2022b). In combination, the bibliographic references used for each entry (Supplementary these features provide abundant information about the palae- Data, Table 1). In order to avoid taxonomic misidentifications, obiogeographic distribution of clavatoraceans in many parts of the only those references with an appropriate description and World over this time interval. illustration of the microfossils were considered. Twenty-six taxa Previous studies on clavatoracean palaeobiogeography have (i.e. 16 species and its varieties) were analysed including species mainly focused on the historical biogeography of a few species that and anagenetic varieties within species, enabling better charac- attained a sub-cosmopolitan to cosmopolitan distributional range, terization of the species’ evolution. allowing for inter-continental biostratigraphic correlations (e.g. Nine palaeogeographic areas were considered: 1) Asian Main- Martín-Closas and Wang, 2008; Martín-Closas, 2015). Such is the land (ASM), mainly including data from China, but also more case of Atopochara trivolvis Peck and Clavator harrisii Peck, which sparse data from Kazakhstan, Uzbekistan, Tajikistan, Mongolia, were cosmopolitan during some stages (e.g. the Aptian), at least in a Korea, and Japan; 2) North European landmass (NE), including wide tropical to paratropical latitudinal belt in the Northern basins from North Germany and the English Wealden; 3) Central Hemisphere. In contrast, little is known about the biogeography of Massif Islands (CM), including data from the islands located south most of the other clavatoracean species, which did not expand over from the Massif, i.e. the Subalpine Chains in eastern France and the large areas of the Cretaceous World but remained confined within Jura Mountains in Switzerland; 4) Iberian Island (IB), including the palaeogeographic domain of the present-day Europe, North data from Spain, Portugal, and Sardinia (Italy); 5) Moesian-Dacian Africa, and the Middle East. In the Early Cretaceous this palae- Island and neighbouring peninsulas (MD), including data from ogeographic domain was covered by a shallow tropical sea, the present Hungary, Romania, Moldova, southeast Ukraine, and Tethys, interspersed with numerous islands of different sizes, southwest Russia; 6) North American Mainland (NAM), including forming what is generally known as the Central Tethyan Archipel- data from the USA (mainly south of the Interior Seaway); 7) South ago (CTA). American Mainland (SAM), including data from Argentina and The aim of the present study was to characterize the biogeo- Brazil; 8) North African islands (NAF), including data from graphic distribution of all clavatoracean species during the Morocco, Algeria, and Tunisia; and finally, 9) Levantine Island and Barremianeearly Aptian, paying special attention to the area northeast Africa (LEA), including Lebanon, Israel, and the eastern where they were more abundant and diverse, the CTA, in order to coast of Africa, mainly Somalia. Some areas were excluded from test the hypothesis that this archipelago was home to a particular the study due to the absence of data. These correspond to northern biota, as suggested earlier for other important groups, such as South America, central and south Africa, and Oceania (Australia dinosaurs (Csiki-Sava et al., 2015). To this end, we selected the and New Zealand). Early Cretaceous, Barremianeearly Aptian (129.4e120 Ma) time interval, which corresponds to the time when the Clavatoraceae 2.2. Data analysis family attained their maximum diversification (Martín-Closas and Serra-Kiel, 1991). This time interval is also well constrained bio- The dataset on clavatoracean occurrences during the stratigraphically since it corresponds to the Eurasian biozones of Barremianeearly Aptian was plotted in a binary matrix, consid- Atopochara trivolvis triquetra, Hemiclavator neimongolensis var. ering the species absence (0) or presence (1) (Table 1). This data neimongolensis, and Clavator gautieri var. jiuquanensis defined by matrix was then studied by cluster analysis using the software Perez-Cano et al. (2022a). The results are intended to provide not PAST 4.03 developed by Hammer et al. (2001), in order to identify only a clear picture of the biogeographic distribution of a fossil and test links between species and their palaeogeographic plant group with an exceptionally rich fossil record (Feist et al., distribution. All cluster analyses used the Pair-Group (UPGMA) 2005), but also to serve as a proxy for comparison with other algorithm, since it favours joining two groups based on the less abundant non-marine groups, such as vascular plants (taxa average distance between all members, which to our from mega-remains), insects, or vertebrates. Furthermore, it will understanding is closer to the purposes of the biogeographic be useful to test current hypotheses about the spatial relationships similarity between areas. More than 20 similarity indexes were between palaeo-landmasses and the effects of climate on very tested and those with the highest correlation coefficients were sensitive environments, such as lakes and ponds. Lastly, the compared before a final dendrogram was selected to represent biogeographic distribution of charophyte species will provide key the results. 2 A. Vicente, J. Sanjuan, J. Perez-Cano et al. Cretaceous Research 154 (2024) 105752 Table 1 Data matrix for the cluster analyses of the Clavatoracean distribution during in the Barremianeearly Aptian. Species/Geographic area MATRIX Worldwide Laurasia Gondwana ASM NE CM IB MD NAM SAM NAF LEA Subfamily Atopocharoidae (Grambast) emend. Martín-Closas ex Schudack Echinochara (Peck) emend. Pe rez-Cano, Bover-Arnal et Martín-Closas Echinochara lazarii (Martín-Closas) Mojon ex Pe rez-Cano, Bover-Arnal et Martín-Closas 0 0 1 1 0 0 1 1 0 Globator Grambast Globator maillardii var. trochilisoides (Grambast) Martín-Closas 0 0 1 1 1 0 0 1 0 Globator maillardii var. biutricularis Vicente et Martín-Closas 0 0 0 1 1 0 0 1 0 Atopochara Peck *Atopochara trivolvis var. triquetra (Grambast) Martín-Closas 1 1 1 1 1 0 1 1 1 *Atopochara trivolvis var. trivolvis Peck 1 1 1 1 1 1 1 1 1 Subfamily Clavatoroidae (Grambast) emend. Martín-Closas ex Schudack Clavator (Reid et Groves) emend. Martín-Closas ex Schudack Clavator grovesii var. gauteri and C. grovesii var. combei (Grambast) Martín-Closas 0 1 1 1 1 0 0 0 0 *Clavator grovesii var. jiuquanensis (Wang) Martín-Closas and C. g. var. jiuquanensis-corrugatus 1 1 1 1 1 0 0 0 0 Clavator ampullaceus var. ampullaceus (Grambast et Lorch) Martín-Closas 0 0 0 0 0 0 0 0 1 Clavator ampullaceus var. latibracteatus Sanjuan, Vicente, Pe rez-Cano, Stoica et Martín-Closas 0 0 0 0 1 0 0 0 0 *Clavator harrisii var. harrisii Peck 1 0 0 1 1 1 1 1 0 Clavator harrisii var. reyi (Grambast-Fessard) Martín-Closas and C. harrisii var. reyi-zavialensis 0 0 1 1 1 0 0 1 0 *Clavator harrisii var. dongjingensis (Hu et Zeng) Martín-Closas 1 0 0 1 0 0 0 0 0 *Clavator calcitrapus var. jiangluonensis (Wang et Li) Pe rez-Cano, Bover-Arnal et Martín-Closas 1 1 0 1 0 0 1 0 0 Clavator calcitrapus var. calcitrapus (Grambast) Martín-Closas ex Schudack 0 0 0 1 0 0 0 0 0 Ascidiella (Grambast) emend. Martín-Closas ex Schudack Ascidiella stellata var. stellata (Martín-Closas et Grambast-Fessard) Martín-Closas ex Schudack 0 0 0 1 0 0 0 0 0 Ascidiella stellata var. lata Martín-Closas 0 0 0 1 0 0 0 0 0 Ascidiella triquetra (Grambast) Martín-Closas ex Schudack 0 0 1 1 0 0 0 0 0 Ascidiella cruciata (Grambast) Martín-Closas ex Schudack 0 0 1 1 1 0 0 0 0 Ascidiella iberica var. iberica (Grambast) Martín-Closas ex Schudack 0 1 0 1 1 0 0 0 0 Ascidiella iberica var. inflata (Grambast-Fessard) Martín-Closas 0 0 0 1 0 0 0 1 0 Ascidiella reticulata var. reticulata Grambast et Lorch 0 0 0 0 0 0 0 0 1 Hemiclavator Wang et Lu Hemiclavator adnatus (Martín-Closas et Grambast-Fessard) Schudack 0 0 0 1 1 0 0 1 0 Hemiclavator neimongolensis var. posticecaptus (Martín-Closas et Grambast-Fessard) Martín-Closas 0 0 1 1 1 0 0 1 0 *Hemiclavator neimongolensis var. neimongolensis Wang et Lu 1 0 1 1 1 0 0 0 0 Pseudoglobator Grambast Pseudoglobator fourcadei Grambast 0 0 0 1 0 0 0 0 0 Pseudoglobator paucibracteatus Martín-Closas et Grambast-Fessard 0 0 1 1 1 0 0 0 0 Abbreviations: ASM, Asian mainland; CM, Central Massif Island; IB, Iberian Island; LEA, Levantine Island and NE Africa; MD, Moesian-Dacian Island and neighbouring pen- insulas; NAF, North African islands; NAM, North America; NE, Northern Europe; SAM, South America. Sub-cosmopolitan to cosmopolitan species are marked with an asterisk (*). 3. Results (Grambast) Martín-Closas, Ascidiella cruciata (Grambast) Martín- Closas ex Schudack, Ascidiella iberica var. inflata (Grambast-Fessard) The Chord similarity index was selected for cluster analysis Martín-Closas, Hemiclavator neimongolensis var. posticecaptus because of its higher cophenetic correlation coefficient (0.9094). (Martín-Closas et Grambast-Fessard) Martín-Closas, and Pseudo- The dendrogram obtained shows three main separated branches globator paucibracteatus Martín-Closas et Grambast-Fessard (Fig. 2). (Fig. 1A). The first of these branches represents the Central Tethyan These taxa represent about one third (ca. 35%) of the 26 clavator- Archipelago (CTA) which is composed of three different subgroups, acean taxa reported worldwide. one containing the islands bordering the Central Massif Island (CM) Within the CTA group, the North African basins are placed in a to the South, another with the central Iberian (IB) and Moesian- different branch (Fig. 1A), mainly due to the absence of some taxa Dacian (MD) islands, and a third group represented by the North (either species or varieties) belonging to genus Ascidiella Grambast African islands (NAF). A second main branch is an heterogenous (e.g. A. triquetra and A. cruciata), Clavator (Reid et Groves) emend. group that includes the neighbouring mainland from Northern Martín-Closas ex Schudack (e.g. C. grovesii gautieri and C. grovesii- Europe (NE) and the continents North America (NAM), South combei) or Pseudoglobator Grambast (e.g. Pseudoglobator pauci- America (SAM), and Asia (ASM). Finally, the third main branch, bracteatus). The islands bordering the Central Massif to the south includes the Levantine Island and NE Africa (LEA), which is (Subalpine Chains and Jura Mountains) are separated due to the completely isolated from the other groups in the dendrogram absence of Globator maillardii var. biutricularis, Hemiclavator adna- (Fig. 1A). tus, and Ascidiella iberica var. inflata. On the other hand, the Iberian The separation of the CTA (CM, IB, MD, and NAF) from all the Island (IB) and the Moesian-Dacian basins (MD) share a large other areas is based on the common occurrence of many clavator- number of taxa (Fig. 1A-B). acean species and varieties unique to the CTA islands (CTA in The second group (NE, ASM, SAM, and NAM) observed in the Fig. 1B). This branch contains the CTA endemic taxa Globator dendrogram (Fig. 1A) is justified by the occurrence of only maillardii var. trochiliscoides (Grambast) Martín-Closas, G. maillardii cosmopolitan and sub-cosmopolitan (Eurasian) species such as var. biutricularis Vicente et Martín-Closas, Clavator harrisii var. reyi Atopochara trivolvis var. triquetra (Grambast), A. trivolvis var. triv- (Grambast-Fessard) Martín-Closas, Hemiclavator adnatus (Martín- olvis Peck, Clavator grovesii var. jiuquanensis (Wang) Martín-Closas, Closas et Grambast-Fessard) Schudack, Ascidiella triquetra Clavator harrisii var. harrisii Peck, and Clavator calcitrapus var. 3 A. Vicente, J. Sanjuan, J. Perez-Cano et al. 4 Cretaceous Research 154 (2024) 105752 Fig. 1. Biogeographic analysis of the Barremianeearly Aptian clavatoraceans in a worldwide context, obtained from a cluster analysis with PAST using the Pair Group algorithm and Chord similarity index. A. Dendrogram of area similarity. B. Dendrogram of species distribution similarity. C. Palaeobiogeographic distribution of the areas represented by the main branches of the dendrogram A on the World palaeomap of the BarremianeAptian boundary (ca. 121.4 Ma). Palaeogeography modified from Martín-Closas and Wang (2008). See acronyms for areas in the Materials and Methods section. A. Vicente, J. Sanjuan, J. Perez-Cano et al. Cretaceous Research 154 (2024) 105752 5 A. Vicente, J. Sanjuan, J. Perez-Cano et al. Cretaceous Research 154 (2024) 105752 jiangluonensis (Wang et Li) Pe rez-Cano, Bover-Arnal et Martín- Riveline et al., 1996; Mojon, 2002; Vicente and Martín-Closas, 2013; Closas. Northern European basins (NE) are ranged in this group of Perez-Cano et al., 2022a). This geological application has yielded regions due to the relative abundance of cosmopolitan and Eurasian numerous records from many basins in the CTA. In the subfamily species although they also contain a few taxa characteristic of the Clavatoroidae two genera, Ascidiella Grambast and Pseudoglobator, CTA; i.e. Clavator grovesii var. combei (Grambast) Martín-Closas and are most typical of the CTA (Figs. 1, 3). The former is a relatively Ascidiella iberica var. iberica Grambast. diverse genus that includes A. stellata, A. triquetra, A. cruciata, and Lastly, the separation of the Levantine Island and surrounding A. iberica in the CTA. However, some species occurred also on areas from all previous groups is well justified by the unique neighbouring coasts. Such is the case of A. reticulata var. reticulata, occurrence in this region of Clavator ampullaceus var. ampullaceus endemic from the LEA or A. iberica var. iberica recorded both in the (Grambast et Lorch) Martín-Closas and Ascidiella reticulata var. CTA and in the North European landmass. In contrast, genus Pseu- reticulata Grambast et Lorch (Fig. 1AeB). doglobator is formed by 2 species, P. paucibracteatus and P. fourcadei, both of which are unique to the CTA. Three more genera, belonging 4. Discussion to the family Clavatoraceae (i.e. Atopochara, Clavator, and Hemi- clavator) were present both within and beyond the CTA, some 4.1. Definition of a charophyte bioprovince in the Central Tethyan species even attaining cosmopolitanism. However, the species from Archipelago these genera appeared first on the islands of the CTA (Martín-Closas and Wang, 2008; Martín-Closas, 2015). This is the case, for instance, The results obtained herein allow confirming the presence of a of Atopochara trivolvis, which appeared in the CTA during the Ber- separate biogeographic region in the Central Tethyan Archipelago riasian, and reach cosmopolitanism in a latitudinal belt of the (CTA). It is characterised by the occurrence of unique and abundant Northern Hemisphere during the Aptian (Martín-Closas and Wang, clavatoracean charophyte species at least during the Barremian and 2008). Clavator calcitrapus was mainly an early Barremian species in early Aptian, but probably also throughout the entire Early Creta- Eurasia and South America (Martín-Closas, 2015), but its first ceous. This region consists of a core of islands formed by Iberia representative, Clavator calcitrapus var. jiangluoensis, occurred in (Spain and Portugal), the basins surrounding the Central Massif the late Hauterivian of the CTA (Feist et al., 1995). In addition, (east France and Switzerland), the Moesian-Dacian platform islands Hemiclavator neimongolensis var. neimongolensis is widely repre- (mainly Bulgaria, Hungary, Romania, Moldova, and parts of sented in Eurasia during the Barremian and early Aptian (Martín- Ukraine), and finally, the North African Islands (mainly Tunisia and Closas, 2015), but it first appears in the CTA at the end of the Ber- Morocco). This region differs from most of the surrounding riasian, where the anagenetic variety Hemiclavator neimongolensis biogeographic regions of the World, including the North and South var. posticecaptus was found on both extremes of the main core American mainlands (NAM and SAM respectively), and the Asian islands, i.e. Spain in the Iberian Island (Martín-Closas, 2000) and Mainland (ASM), because these regions only contain clavatoracean Romania in the Moesian-Dacian Island (Sanjuan et al., 2021a). species considered sub-cosmopolitan or cosmopolitan (Fig. 1). Of Beyond the CTA the Barremianeearly Aptian charophyte floras particular note among the regions within this group is the North were dominated by two other families, the family Characeae, rep- European landmass (NE), including English and German basins, resented mainly by the genera Aclistochara (Peck) Schudack, Mes- because it has transitional floras between the CTA and its neigh- ochara Grambast, and Sphaerochara (Ma €dler) Soulie -Ma€rsche, and bouring continents. Finally, the Levantine and NE African (LEA) the family Porocharaceae (Grambast) emend. Schudack, charac- flora shows an affinity with the CTA in being dominated by clav- terized mainly by the genera Porochara Grambast and Feistiella atoraceans, but also fundamentally differs from them in hosting a Schudack, only associated with the cosmopolitan or sub- significant proportion of its own endemic taxa. This region forms an cosmopolitan (i.e. Eurasian) representatives of the Clavatoraceae isolated branch of the dendrogram obtained (Fig. 1A), which may (Martín-Closas and Wang, 2008; Martín-Closas, 2015). Some ex- represent a different bioprovince from the CTA, perhaps containing amples of this type of floras have recently been described in China the equatorial charophytes of the Early Cretaceous. However, the (Li et al., 2020) and in South America (De Sosa Toma s et al., 2022) absence of data from Central Africa and the north of South America, for the interval studied (Barremianeearly Aptian). which formed a single landmass during the Barremian, prevents an In sum, all the available data suggest that a particular char- assessment of this hypothesis now. ophyte bioprovince can be proposed for the Barremianeearly Within the family Clavatoraceae, which was composed of ca. 26 Aptian Central Tethyan Archipelago (CTA), and that it probably evolutionary species from the Late Jurassic until the latest Creta- persisted throughout the entire Early Cretaceous. Based on the ceous according to Martín-Closas (1996), at least 2 clavatoracean abundance and biodiversity of clavatoracean charophytes endemic genera (i.e. Pseudoglobator and Globator Grambast) and up to 8 to this archipelago, this bioprovince can be called the “Clavator- species (Table 1; Fig. 1B) belonging to two different subfamilies (i.e. acean Bioprovince”. Atopocharoidae (Grambast) emend. Martín-Closas ex Schudack and Clavatoroidae (Grambast) emend. Martín-Closas ex Schudack) are 4.2. Climatic constraints of the Clavatoracean Bioprovince unique to the CTA during the Barremianeearly Aptian. In the sub- family Atopocharoidae the monospecific genus Globator, with the The results obtained reveal that the Clavatoracean Bioprovince species Globator maillardii (Saporta), has been reported only from is separated from neighbouring areas by latitudinal boundaries. The the CTA (Figs. 1, 3). This species persists from the Tithonian until the northern boundary is represented by the North European land- early Aptian, evolving into an anagenetic lineage that has been used mass, which is located at a palaeo-latitude of ca. 35e45 N (Fig. 3). intensively for biostratigraphic purposes (e.g. Grambast, 1974; The main core of islands of the CTA ranges in latitude from 10 to Fig. 2. Utricles of clavatoracean species characteristic of the Central Tethyan Archipelago. AeB. Globator maillardii var. trochiliscoides (Southern Dobrogea, Romania; from Sanjuan et al., 2021a). A. Lateral view; B. basal view. CeD. Globator maillardii var. biutricularis (Southern Dobrogea, Romania; from Sanjuan et al., 2021a, b). C. Lateral view; D. Basal view. EeF. Clavator grovesii var. gautieri (Iberian Chain; Spain; from Perez-Cano et al., 2022a). E. Lateral view; F. Adaxial view. GeH. Ascidiella triquetra (Iberian Chain, Spain; from Pe rez-Cano et al., 2020). G. Oblique-abaxial view; H. Apical view. IeJ. Ascidiella cruciata (Iberian Chain, Spain; from Vicente and Martín-Closas, 2013). I. Lateral view; J. Apical view. K. Ascidiella iberica var. inflata (lateral view; Iberian Chain, Spain). LeM. Hemiclavator adnatus (Iberian Chain; Spain; from Pe rez-Cano et al., 2020). L. Abaxial view; M. Lateral view. NeO. Hemiclavator neimongolensis var. posticecaptus (Iberian Chain, Spain; from Pe rez-Cano et al., 2020). N. Abaxial view. O. Lateral view. PeQ. Pseudoglobator paucibracteatus (Subalpine Chains, France; from Martín-Closas et al., 2009). P. Lateral view; Q. Basal view. 6 A. Vicente, J. Sanjuan, J. Perez-Cano et al. Cretaceous Research 154 (2024) 105752 Fig. 3. Distribution of species characteristic of the Central Tethyan Archipelago during the Barremian and early Aptian. Palaeogeographic map modified from http://deeptimemaps. com (accessed November 2022) and Dercourt et al. (2000). 35 N (Fig. 3). South of this area, the Levantine Island and the NE belt would include lakes and wetlands with relatively high tem- African region would have belonged to a tropical-equatorial belt peratures throughout the year but with high seasonal changes in ranging from ca. 10 N to about 10 S (Fig. 3). The separation be- precipitation (monsoonal climates), while the tropical belt would tween these three latitudinal belts suggests that the charophyte have been influenced by high temperatures and precipitation but floras were climatically controlled. As a matter of fact, they corre- with low seasonal changes (Scotese, 2003). spond quite precisely with the temperate, paratropical, and tropical Besides the more evident latitudinal distribution of the CTA, climatic belts defined for the Early Cretaceous by Scotese (2003). there were also longitudinal relationships between them. This is The islands in the temperate belt would have been influenced by observed in the occurrence of common taxa in the same longitu- wide temperature ranges and high seasonality. The paratropical dinal strip, i.e. separately in the eastern and the western islands of 7 A. Vicente, J. Sanjuan, J. Perez-Cano et al. Cretaceous Research 154 (2024) 105752 the archipelago. In the eastern part of the archipelago Clavator least since the MiddleeLate Jurassic, being consequently one of the ampullaceus is shared between the Levantine-NE African area and most useful groups in biostratigraphy, stratigraphic correlation, the Moesian-Dacian area, i.e. across the paratropical and tropical palaeoecology, and palaeobiogeography of the aquatic non-marine belts (Fig. 3). However, this species developed two different vari- domain. eties in each area, respectively; Clavator ampullaceus var. ampulla- Mesozoic non-marine ostracods belong to the three superfamilies ceus in the South (Sanjuan et al., 2021b and references therein) and Cypridoidea Baird, Cytheroidea Baird, and Darwinuloidea Brady and C. ampullaceus var. latibracteatus in the North (Sanjuan et al., Norman, among which the first has experienced a particular 2021a). In the two areas these varieties grew in euryhaline to ‘explosive’ radiation during the Late JurassiceEarly Cretaceous freshwater wetlands. The same palaeogeographic pattern can be worldwide (Whatley, 1990). Species of the two genera Cypridea observed in the western islands, for instance in species such as (Cypridoidea) Bosquet and Theriosynoecum Branson (Cytheroidea) Ascidiella iberica, which occurs only in the North African basins, are the main tools used in biostratigraphy of the Purbeck-Wealden Iberian basins, and British Island basins, i.e. across the paratropical facies (Horne, 2009) and a large amount of distribution data are and temperate belts and also in similar facies (mainly freshwater available about them during the Barremianeearly Aptian interval lakes). This north-south distribution pattern appears to be limited studied here. to a few species and might be related to specific dispersal mecha- The Wealden Group, as defined in its English type locality, is nisms. Nowadays some charophytes are dispersed over long- chronostratigraphically calibrated to the Hauterivianeearly Aptian distances in the digestive tract of migratory birds (Proctor, 1980), time interval, assigned to the ostracod biozone Theriosynoecum which may also have been the case in the early Aptian, by which fittoni of Horne (2009). The upper Wealden, Barremianeearly time aquatic birds with well-developed sternums, capable of un- Aptian in age, is assigned to the subzone Cypridea fasciata of dertaking long-duration flights already existed (You et al., 2006). Horne (2009), and mainly characterized by species of genus Cyp- ridea such as Cypridea tenuis Anderson, C. fasciata Anderson, C. 4.3. Endemism within large islands clavata Anderson, C. comptonensis Anderson, C. bogdenensis Anderson, C. brendae Anderson, C. marina Anderson, C. pseudo- The Iberian Island was one of the largest islands in the Central marina Anderson, C. insulae Anderson, C. recta Anderson, C. hispida Tethyan Archipelago and is probably the region for which most data Anderson, C. pumila Anderson, C. dorsospinata Anderson, C. val- are available on Early Cretaceous charophytes of the CTA. It appears densis Sowerby, and C. tuberculata Sowerby. They are usually that this island was sufficiently large and isolated to give rise to associated to Theriosynoecum fittoni Mantell, Rhinocypris jurassica endemism (Figs. 1 and 3). Such is the case of Pseudoglobator four- Martin, Mantelliana mantelli Jones, Fabanella boloniensis Jones, and cadei (Fig. 3), which is only known from the Prebetic Chain (Martín- Alicenula leguminella Forbes (Anderson, 1985; Horne, 1995, 2009). Closas, 2000). Apart from this endemism, Iberia also had other Such a species assemblage has also been described from Northwest possible Barremian endemic taxa, such as Clavator calcitrapus var. Germany (Martin, 1940; Wolburg, 1959; Schudack, 1994; Elstner rez-Cano et al., 2020). Endemic calcitrapus or Ascidiella stellata (Pe and Mutterlose, 1997) and Denmark (Christensen, 1963), within species may also have occurred in other large islands of the CTA. the North European landmass. Some Wealden ostracod species This is possibly the case of Clavator ampullaceus var. latibracteatus were also reported from southern European basins, mainly from described from Romania, on the Moesian-Dacian Island, by Sanjuan the Iberian Island (IB), i.e. from Spain (Kneuper-Haack, 1966; et al. (2021a). However, in spite of these possible single-island Brenner, 1976; Swain, 1993; Schudack and Schudack, 2009) and endemisms, the homogeneity of the CTA bioprovince in terms of Portugal (Damotte et al., 1990; Cabral, 1995), as well as from the taxonomy was high throughout the entire time interval studied. North African domain, i.e. Tunisia (Trabelsi et al., 2015, 2021; Frequent sea-level changes during the Barremian and early Aptian Chnayna et al., 2021) and Morocco (Andreu, 1989, 1991; Andreu probably helped to facilitate floral exchange between neighbouring et al., 2003). Differences in the distribution of some Wealden islands and to maintain the biogeographic identity of the archi- ostracod assemblages highlights a north to south distribution pelago over a long period of time. pattern. These data allow for a biogeographic comparison with the re- 4.4. Comparisons with the biogeographic distribution of other non- sults presented here from charophytes. First the extension of the marine fossil groups Clavatoracean Bioprovince coincides broadly with the distribution pattern of ostracods in the same time interval. The northern Palaeobiogeographic zonation has a long tradition in palae- boundary of the Clavatoracean bioprovince is represented in the obotanical studies, mainly in palynology. The proposals available to ostracod faunas by a latitudinal boundary between the northern date of a floristic zonation for the Early Cretaceous are based in Wealden assemblages (English, German, and Danish faunas) and sporomorphs and result in the definition of very large floristic those from the south (Iberia and North Africa), the latter corre- provinces, which are approximately equivalent to latitudinal strips sponding to the Tethyan islands of the CTA. The southern boundary (e.g. Herngreen and Chlonova, 1981; Herngreen et al., 1996). This is of the Clavatoracean Bioprovince, placed at 10 N latitude, is much possibly due to a mainly climatically controlled distribution of the better defined in ostracod faunas. Barremianelower Aptian non- sporomorph megaproducers used for biogeographic zonation, most marine ostracod assemblages from the Levantine domain, mainly of them from anemophilous (i.e. wind pollinated) plants. Wind- from Israel (Rosenfeld and Raab, 1984) and Lebanon (Bischoff, 1963) dispersal of sporomorphs facilitates their use in large-scale are formed by species completely different from the coeval Weal- biostratigraphic correlation (e.g. Ied and Tahoun, 2019), but un- den fauna, hosting a significant proportion of endemic cypridean dermines any possible comparison with the results obtained here species, such as Cypridea libanensis Bischoff, C. heliopolisensis Bis- from charophytes. Other terrestrial groups, such as vertebrates and choff, C. aaleyensis Bischoff, C. baidarensis Bischoff, C. quadrangulata insects provide valuable biogeographic data for comparison (e.g. Brenner, C. orientalis Bischoff, and C. sidonensis Bischoff. From these Peris et al., 2016), yet insufficient to undertake a biogeographic Levantine species, C. heliopolisensis has been recovered also from zonation. In order to test these results with other organisms that Brazil (Kro€ mmelbein and Weber, 1971; Beurlen et al., 1994), and C. have fossil records and palaeoenvironmental conditions compara- aaleyensis from North Somalia (Luger and Schudack, 2001). This ble with charophytes, we chosed non-marine ostracods. They are suggests a southward extension and biogeographic connection of among the most common fossils in aquatic continental settings, at these faunas across the Gondwanan landmass. These differences 8 A. Vicente, J. Sanjuan, J. Perez-Cano et al. Cretaceous Research 154 (2024) 105752 between the biogeographic distribution of the Wealden ostracod distribution. The Characeae and the paraphyletic group of the species in the north and the Gondwanan faunas in the south Porocharaceae are the two charophyte families dominating outside highlights, also in ostracods, a climatic control during the late Early the Central Tethyan Archipelago. Cretaceous, as was already suggested by Anderson (1973). The CTA clavatoracean bioprovince shows a latitudinal pattern Second, the dispersal of ostracods between islands of the CTA which is certainly related to climatic constraints. However lon- appears to have been also common fact, like in charophytes. For gitudinal strips of regions within the bioprovince and the instance, cypridean species such as Cypridea cyrtorostralis Brenner neighbouring areas suggest that other factors may have impinged and C. ventriosa Brenner, though to be endemic from Iberia, were the distribution of charophyte floras within the bioprovince. In recently reported from the Central Tunisian Atlas (Trabelsi et al., the east of the CTA this is the case of the links between Moesia- 2015, 2021; Chnayna et al., 2021). Dispersal of Wealden ostracod Dacia and the putative equatorial bioprovince, while to the west species within CTA is also confirmed by the occurrence of the of the archipelago, North Africa and Iberia shared some species traditionally West-European species Theriosynoecum fittoni, Cypri- with the North European landmass. These longitudinal affinities dea recta, and C. fasciata in Romania, i.e. in the Moesian-Dacian may be linked to the dispersal vectors of particular species that domain (Antoniade, 2016). migrated in a north to south axis. Other possible variables in the Another important similitude in the biogeography of biogeographic distribution of clavatoraceans include the palae- Barremianeearly Aptian charophytes and non-marine ostracods is oecological constraints of the different species. They are of that some of them, notably genera Cypridea, Harbinia Tsao, Ther- difficult evaluation at the present state of the art, since only a few iosynoecum, and Alicenula Rossetti et Martens, achieved cosmopol- palaeoecological studies have been carried out to date with itan range during this time interval. They have been described from clavatoraceans, and most of them were devoted to the Iberian Europe and North America (Sames, 2011a, b and references therein), floras. North Africa (Andreu, 1991; Trabelsi et al., 2015; Chnayna et al., In spite of all these superposed climatic and biogeographic 2021), South America and West Africa (Grekoff and Kro € mmelbein, controls, the homogeneity of the CTA charophyte bioprovince 1967; Poropat and Colin, 2012 and references therein), East Africa remained high during the time interval studied, i.e. several millions (Luger and Schudack, 2001; Sames, 2008), Asia (Hou et al., 2002; of years, as attested by the low number of endemic species and the Hayashi, 2006), Middle East (Rosenfeld and Raab, 1984; Sanjuan restriction of regional differences to the infraspecific rank. Similar et al., 2021b). Accordingly, Trabelsi et al. (2015), and Sames and biogeographic patterns are found for non-marine ostracods sug- Trabelsi (2018) suggested that peri-Tethyan islands could have gesting that the CTA charophyte bioprovince can be extrapolated at worked as effective bridges for non-marine ostracods dispersal by least to other aquatic non-marine benthic groups, while data ‘island-hopping’ of larger animals. available from terrestrial groups are either too limited (e.g. verte- brates and arthropods) or provide a very different biogeographic 5. Conclusions signal (sporomorphs) due to very distinct dispersal vectors. Charophyte bioprovincialism is first explored in the charophyte Data availability fossil record. Previous attempts to study the biogeographic distri- bution of fossil charophytes were limited to describing the Data will be made available on request. composition of charophyte floras in the different continents along geological time, without providing a biogeographic analysis of Acknowledgements bioprovincialism. The Central Tethyan Archipelago (CTA) is proposed here to host This study was funded by and contributes to the research project a particular subtropical bioprovince at least during the IBERINSULA, Grant PID 2020-113912GB-100 funded by MCIN/AEI/ Barremianeearly Aptian interval. The new bioprovince is charac- 10.13039/501100011033 (Spanish Ministry of Science and Innova- terized by up to 2 genera and 8 species belonging to the family tion, Spanish Research Agency), and to project “Geologia Sed- Clavatoraceae, which are unique to this area, i.e. most of the imentaria” (2022 SGR 00349), funded by AGAUR (Generalitat de Tethyan Europe and North Africa. The Middle East and north-east Catalunya, Catalan Autonomous Government). AV's postdoctoral Africa had evident relationships with the CTA charophyte bio- research was funded by the project A1S19598 of the Consejo province, especially because their charophyte floras were also Nacional de Ciencia y Tecnología (CONAHCyT) from Mexico. JP-C dominated by the same family, but with a different taxonomic Margaria Salas postdoctoral contract and research was supported composition at the species level. Although the data about this latter by the Ministerio de Universidades of Spain, “Plan de recuperacio n, area are insufficient yet, a future definition of a different, properly transformacio n y resiliencia” of the Spanish Government and the equatorial bioprovince for Early Cretaceous charophytes in this Next Generation founds from the European Union. KT contribution southern domain of the Tethys would seem justified, and is already is part of the IGCP Project 710-Western Tethys meets Eastern clear for other aquatic groups, such as ostracods. Tethys, subproject “Non-marine e marine correlation and sea-level Outside the Central Tethyan Archipelago charophyte bio- changes in the Mid-Jurassic Tethys: Tectonic versus climate events” provincialism during the Early Cretaceous remains undefined. € funded by the Austrian Academy of Science (OAW). The English text However, bioprovinces in these areas will be possibly based in has been corrected by Dr. Michael Maudsley (Universitat de Bar- charophyte families different than the Clavatoraceae, since only a celona). We would like to thank the editor Eduardo Koutsoukos and few species of this family migrated from the CTA to the neigh- an anonymous reviewer who provided valuable comments during bouring continents to reach a sub-cosmopolitan or cosmopolitan the peer-review process. 9 A. 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Ostracods as Proxies for Marginal Marine to Non-Marine Intervals in the Mid-Cretaceous Carbonate Platform of the Central Tunisian Atlas (North Africa) Response to Major Short-Term Sea-Level Falls. Cretaceous Research 117, 104581. Appendix A. Supplementary data Vicente, A., Martín-Closas, C., 2013. Lower Cretaceous charophytes from the Ser- ranía de Cuenca, Iberian chain: Taxonomy, biostratigraphy and palaeoecology. Supplementary data to this article can be found online at https://doi.org/10. Cretaceous Research 40, 227e242. 1016/j.cretres.2023.105752. 11