Algal diversity during the onset of the Late Palaeozoic Ice Age in low-latitude basins of the Western Palaeotethys

Earth-Science Reviews 246 (2023) 104596 Available online 14 October 2023 0012-8252/© 2023 The Author(s). Published by Elsevier B.V. This is an open access article under the CC BY-NC license (http://creativecommons.org/licenses/bync/4.0/). Algal diversity during the onset of the Late Paleozoic Ice Age in low-latitude basins of the Western Palaeotethys Pedro C´ ozar a , b , * , Ian D. Somerville c , Mark W. Hounslow d , e , J. Ricardo Mateos-Carralafuente b , Ismael Coronado f a Instituto de Geociencias (CSIC-UCM), c/ Severo Ochoa 7, 28040 Madrid, Spain b Departamento de Geodin´ amica, Estratigrafía y Paleontología, Facultad de Ciencias Geol´ ogicas, Universidad Complutense de Madrid, c/ Jos´ e Antonio Novais, 12, 28040 Madrid, Spain c UCD School of Earth Sciences, University College Dublin, Belfield, Dublin 4, Ireland d Lancaster Environment Centre, Lancaster University, Lancaster LA1 4YW, UK e Earth, Ocean and Ecological Sciences, University of Liverpool, Jane Herdman Building, Liverpool L69 3GP, UK f Facultad de Ciencias Biol´ ogicas y Ambientales, Universidad de Le´ on, Campus de Vegazana s/n, 24071 Le´ on, Spain ARTICLE INFO Keywords: Calcareous algae Diversity Icehouse LPIA Mississippian Palaeotethys ABSTRACT A study of calcareous algae from the late Vis´ ean to Serpukhovian interval in basins from the western margin of Palaeotethys (basins located currently in Western Europe and Northern Africa) shows varied responses in terms of palaeoecological diversity, specimen abundance, genus richness and taxonomic distinctness for the lowlatitude basins. Algal associations are more similar in cratonic areas, although many local diversity peaks correspond to ecological/environmental factors. Terranes with Laurussian affinities contain the most unusual abundances and seem to be the most affected by palaeogeographic reorganization and local tectonics. Algae from the palaeo-equatorial zone show more dramatic changes than those in the southerly tropics. The varied results suggest that it is not suitable to extrapolate a particular diversity result from a single region to global conditions, since many factors have influenced the regional assemblages. The main taxonomic turnover of most fossil groups occurred from the uppermost Serpukhovian, linked to a severe cooling phase inferred by δ 18 O data from brachiopods. It coincides with the post-glacial phase C1 defined in eastern Australian basins. Most ecological changes occurred from the early Serpukhovian, predating the glacial phase C1 and coinciding with the Main Eustatic Signal. Far-field proxies used to define the onset of icehouse intervals in the LPIA are inconsistent and are located several million years before the main biota changes. This questions the definition of the onset of sustained icehouse glaciation at different levels of the low palaeolatitude Vis´ ean, without the resulting modifications in the warm-water biota. 1. Introduction The Late Paleozoic Ice Age (LPIA) is the longest-lived glaciation recorded in the entire Phanerozoic, and its duration at ~70 Myr, is commonly considered to extend from the Famennian (Late Devonian) to the Sakmarian (Early Permian) (e.g., Monta˜ nez, 2021). Ice accumulation becomes more significant from the middle Vis´ ean onwards, a period where sustained glacial conditions have been inferred (Monta˜ nez, 2021). However, some authors consider the onset of sustained cooling conditions began later, in the Serpukhovian (Fielding et al., 2008a, 2008b, 2023; Ahern and Fielding, 2021). Most glaciogenic signatures are recorded in high-latitude basins (e.g., Fielding et al., 2008a, 2008b, 2023; Limarino et al., 2014; Isbell et al., 2021; L´ opez-Gamundi et al., 2021), whereas in low-latitude areas, the effects of glaciation are archived in stratigraphic architectures and stratal stacking patterns, which are recognised from the Asbian onwards (e.g., Rygel et al., 2008). LPIA-focused studies have generally analysed either the long-term evolution during the Phanerozoic of invertebrate and microfossil groups, such as the brachiopods, rugose corals, conodonts, foraminifers and crinoids (e.g., Sepkoski, 1996; Stanley and Powell, 2003; Alroy et al., 2008; Fan et al., 2020), or alternatively, analysed macroevolutionary processes for shorter periods of time (e.g., Saunders and * Corresponding author at: Instituto de Geociencias (CSIC-UCM), c/ Severo Ochoa 7, 28040 Madrid, Spain. E-mail addresses: [email protected] (P. C´ ozar), [email protected] (I.D. Somerville), [email protected] (M.W. Hounslow), josericm@ucm. es (J.R. Mateos-Carralafuente), [email protected] (I. Coronado). Contents lists available at ScienceDirect Earth-Science Reviews journal homepage: www.elsevier.com/locate/earscirev https://doi.org/10.1016/j.earscirev.2023.104596 Received 21 March 2023; Received in revised form 4 October 2023; Accepted 10 October 2023 Earth-Science Reviews 246 (2023) 104596 2 Ramsbottom, 1986; Raymond et al., 1990; Shen et al., 2006; Wang et al., 2006; Groves and Lee, 2008; Powell, 2008; Groves and Wang, 2009; Segessenman and Kammer, 2018). However, in these studies, a key uncertainty is the chronostratigraphic control on the stratigraphic record, which is not always as precise as would be desirable, leading to differing inferred synchroneity of faunal evolution (Yao et al., 2020; Shi et al., 2021). A second important confounding factor is that biota sensitivity also changes between organisms. Nektonic faunas (such as ammonoids and conodonts) are typically widespread in open-marine conditions in the Carboniferous, but are less sensitive to rapid environmental changes, and consequently were less affected, because they were less impacted by the areal reduction in the continental shelves due to sea-level falls (Brenchley and Newall, 1984). Benthic organisms (such as brachiopods and crinoids) are rather tolerant organisms, and so are widespread from warm-tropical to temperate platforms, as shown for example by modern brachiopods (Giles, 2012). Similarly, Mississippian foraminifers occur in variable types of platforms and conditions, mostly as endobenthic organisms living in the sediment or seawater/sediment interface, and in contrast with other foraminifers, widely accepted as not being associated with endosymbiotic algae (Vachard et al., 2010). Modern calcareous algae and corals, on the other hand, are the most sensitive organisms to stressed conditions in platform environments (e.g., Graham and Wilcox, 2000; Hoegh-Guldberg et al., 2017). Global studies on the taxonomic biodiversity of marine invertebrates (as well as others focused on rugose corals), that include this latter clade have rather divergent results (for example compare Wang et al., 2006, Yao et al., 2020 and Shi et al., 2021). 1.1. Mississippian calcareous algae Studies dedicated to Mississippian calcareous algae are scarce compared to the above-mentioned invertebrate and microfossil groups, with studies mostly focused on algal taxonomy, biostratigraphy and stratigraphic evolution, with less on their palaeoecology and habitat preference. Studies establishing the relationship between ecology and environments during the Mississippian, are generally focused on regions (e.g., Horbury and Adams, 1996; Pille, 2008) or particular environments (e.g., Adams, 1984; C´ ozar et al., 2019). However, only a few Late Paleozoic taxa extend into the Mesozoic and Cenozoic, and hence, comparisons with modern life styles and environmental conditions is not plausible in extinct taxa without modern equivalents (e.g., Algospongia). Mississippian taxa also seem to have survived in distinctly different environments to their modern counterparts (e.g., C´ ozar et al., 2018b). Also in modern carbonate platforms, living bryopsidales contribute three times more fine-grained carbonate sediment than living dasycladales, whereas in sandand gravel-sized deposits, remains of bryopsidales can reach up to 50% in carbonates (Granier, 2012). The genera-balance during the Mississippian is distinctly different, with only some ancestral rhodophytes, scarce bryopsidales and common dasycladales and problematic algae. This scarcity of Mississippian algal studies is a clear gap in knowledge, compared with their importance as ecological indicators in Modern oceans and ancient environments, where they are commonly used to determine important parameters, such as water temperature, bathymetry, salinity and acidification (e.g., Johnson et al., 2014; Leaf et al., 2020). Considering this good potential as ecological indicators, surprisingly little is known about the biodiversity of calcareous algae in relation to the onset of the LPIA, which this study aims to address. The aims are to: (i) characterise the diversity of calcareous algae in lowlatitude basins in the Western Palaeotethys during the late Vis´ ean to Serpukhovian interval. Our dataset includes Carboniferous basins in Western Europe and Northern Africa (including data from northern England, Ireland, Sierra Morena (SW Spain), Cantabrian Zone (N Spain), Moroccan Meseta and Tindouf (S Morocco-W Algeria). These basins were located in the western end of the Palaeotethys, between the equator and southern tropical zones (Fig. 1) and preceded the Gondwana-Laurussia collision, which lead to emergent land for most of the Pennsylvanian in the region (e.g., Nance et al., 2010); (ii) assess critical regional and global factors controlling their diversity; and (iii) contrast the changes in diversity of calcareous algae with those described for other faunas during the LPIA interval. 2. Methodology In this work, bryopsidales, dasycladales, blue-green algae (also known as cyanobacteria) and problematic algae Algospongia are hereafter considered as calcareous algae (Wray, 1977; Pille, 2008; Leaf et al., 2020). The overall taxonomic classifications followed here are those summarised in Pille (2008), and Vachard and C´ ozar (2010) for the problematic algae Algospongia, which in other cases have been interpreted as red algae, dasycladales, bryopsidales or even, foraminifers. Details of the considered algae, as well as their suprageneric classification are included in the Supplementary Information. Data for some calcareous algae used here have not been previously published, such as those of the Tindouf Basin and most of the succession from northern England sections (see Supplementary Information and Appendix A). Data is based on analysis of material from 1977 stratigraphic levels sampled in carbonates and examination of >5100 thin sections. Shaley intervals are omitted for the diversity analysis due to their minor influence on the total diversity recorded for each interval. Genus richness (GR), based on the occurrence of genera compiled from the literature (including originations and extinctions), has the handicap that there are many endemic algal genera (C´ ozar and Somerville, 2021c), which have a wide geographic distribution, but not sufficiently widespread to be considered cosmopolitan for the large oceans (Appendix C in the Supplementary Information). A second handicap is the preferred suprageneric classification, which, depending on the authors, can be considered as valid or not for the same taxa. Hence, to consider only originations/extinctions for substages, is not the best or only approach to determine the actual algal contribution to the geological record for this interval. Furthermore, due to the scarcity of studies of algae, the first appearance and distribution of many algal genera remains questionable (e.g., Mamet, 1991; Pille, 2008; Vachard and C´ ozar, 2010). Any analysis at species or genus levels assessing the diversity of most taxa is questionable, due to data scarcity. Consequently, genera have been grouped into tribes or orders, depending on the overall palaeoecological conditions and only two rather abundant genera (Koninckopora and Ungdarella) have been studied separately. In total, ten groups have been utilised: blue-greens, bryopsidales, dasycladales, rhodophyta, aoujgaliales, calcifoliids, donezellids, moravamminids, as well as the genera Koninckopora and Ungdarella (see Fig. 1. Palaeogeographic map of the studied sections during the Mississippian. 1, Tindouf (southern Morocco); 2, Western Moroccan Meseta; 3, Sierra Morena (SW Spain); 4, Cantabrian Mountains (northern Spain); 5, Montagne Noire (southern France); 6, northern England; 7, Ireland. RPG =Rheic-Palaeotethys gateway, LPT =Laurussian Palaeotethyan terranes, GPT =Gondwana Palaeotethyan terranes. Modified from Blakey (2008). P. C´ ozar et al. Earth-Science Reviews 246 (2023) 104596 3 Supplementary Information, Figs. S1–S5). The diversity of these groups has been analysed independently, as well as the total diversity, abundance and volume, as well as evaluation of various statistical metrics outlined in Table 1. Analysis of the abundance of species and genera was undertaken using sub-division into four classes, rather than actual counts. This emphasises the role of the “rare” species/genera and minimises the importance of the “common” species/genera. For the more unusual algae, the counts for each class are: 1 =1–2 specimens, 2 =3–5 specimens, 3 =6–10 specimens, and 4 ≥10 specimens. For the more common genera (i.e., Koninckopora, donezellids and Ungdarella), these counts are modified, where 3 =6–20 specimens and 4 ≥20 specimens. 3. Stratigraphic successions investigated All the analysed basins are located in the far western part of the Palaeotethys region during the late Vis´ ean to Serpukhovian (and the earliest Bashkirian in some sections where limestones are recorded). These basins occur in stable cratonic areas and mobile terranes involved in tectonics of the Variscan Orogeny and closure of the RheicPalaeotethys gateway (RPG) (e.g., Nance et al., 2010) (Fig. 1). Sections selected include those from northern England and platforms of the Dublin Basin (Ireland) for the Laurussia Platform. As an example of a Gondwana platform succession, that from the northwestern Tindouf Basin in southern Morocco, is included (Fig. 2). In between these two regions, are basins located in the Western Palaeotethys and in western positions closer to the RPG - the Laurussian Palaeotethyan terranes (abbreviated as LPT). These show a higher palaeobiogeographical affinity with Laurussia. In contrast, those in the more eastern positions of the Palaeotethys - Gondwana Palaeotethyan terranes (GPT) are palaeobiogeographically closer to Gondwana, and are located in more open water conditions in the Palaeotethys (C´ ozar et al., 2018a; C´ ozar and Somerville, 2021c). Successions located in the LPT include those from Adarouch (Western Moroccan Meseta) and the Guadiato-Guadalmellato valleys (Sierra Morena in SW Spain). Successions located in the GPT are exemplified by sections in the Montagne Noire (southern France) and the Valdediezma Platform of the Cantabrian Mountains (NW Spain). Sampling in each basin, including the stratigraphic sections, location and biostratigraphic details are included in the Supplementary Information and Appendix A. In addition, the complete succession from Britain and Ireland for the Vis´ ean to Serpukhovian has also been studied, as detailed in Appendix B. Following palaeogeographic reconstructions, such as Blakey (2008), northern England and Ireland were located in equatorial positions at 10◦S, with the LPT and the GPT at about 20–25◦S. The LPT were in more western locations than the GPT, along the closing Rheic-Palaeotethys corridor, and the Tindouf Basin was at about 30◦S, within more tropical and subtropical latitudes (Fig. 1). The timescale used here for the correlation between sections, involves the Vis´ ean Series for the British substages, up to and including the early Brigantian, whereas above this level, the substages of the East European Platform of Russia (EEP) (within the Serpukhovian Series) are used (see shaded parts in Fig. 3). Significantly, in the scale used here, there is no late Brigantian, because, as demonstrated by C´ ozar and Somerville (2014, 2016), it is virtually equivalent to the early Serpukhovian. The precision of the correlation between the Russian and British substages has been subsequently discussed in C´ ozar and Somerville (2021a, 2021b). 4. Palaeoecological diversity (PD) 4.1. Laurussia (northern England and Ireland) As is well known (e.g., Walkden, 1987; Wright and Vanstone, 2001), the late Asbian platform successions in northern England shows marked cyclicity punctuated by emergent surfaces in largely carbonate successions. From the early Brigantian, a second style of cyclicity (‘Yoredaletype’) is developed, composed of thick siliciclastic deposits above carbonates which occur lower in the cycles, usually with coal in the upper part (Fig. 2). These limestone-clastic cycles have been interpreted as the typical stratigraphic pattern of glaciogenic periods, as recognised in the USA and Russia. In general, in northern England, the algal group diversity (D group ) decreases into the Serpukhovian, although some taxa represent the most abundant algae during the Asbian (Table 2; see Supplementary Information Fig. S1A). Nevertheless, the diversity of each particular group allows recognition of four main algal associations (1–4; Supplementary Information Fig. S2). Furthermore, Association-1 can be also subdivided into 1A, 1B and 1C due to differences in abundances, thus making a total of six algal associations, which are variably recognised in time in the different basins (Figs. 4–5). The paucity of algae in the uppermost early Serpukhovian and late Serpukhovian does not permit the recognition of the predominance of any taxa or group. In northern England, Association-1 is commonly recorded in dark grey bioclastic limestones (mostly packstone), which are non-cyclic during the early Asbian but cyclic for the late Asbian (Fig. 2). The three sub-associations (1A, 1B and 1C), show a progressive increase in Ungdarella and progressive reduction in donezellids, whilst the abundance in Koninckopora remains rather constant (Figs. 4A, 6A–6D). Close to the Asbian/Brigantian boundary, there are frequent dark grey to black limestones, composed of dominantly mudstone and wackestone (Fig. 2), commonly interrupted by palaeokarsts (of short duration), with common quiet-water lagoonal facies. Most of the abundant algae in the underlying Asbian become rare or have disappeared, and a bloom of the dasycladales is observed, forming Association-2 (or the dasycladales association; Figs. 4A, 6E). In the middle part of the early Brigantian, there was a dominance of pale grey limestones, and the presence of Falsocalcifolium allows recognition of Association-3 (the Falsocalcifolium association; Figs. 4A, 6F). This interval is also characterised by a reduction in the number of dasycladales. Association-3 is dominant up to the middle part of the early Serpukhovian, whereas, above this level, the occurrence of Calcifolium distinguishes Association-4 (the Calcifolium association; Figs. 4A, 6G). The accompanying algae are similar to those in Association-3. Although the highest D group are observed in the Asbian, the highest peaks of total diversity (D total ) are observed in the early Brigantian (Fig. 4A). The total diversity in the late Serpukhovian is the lowest throughout the Asbian-Serpukhovian interval, since the calcareous algae are rather scarce (Fig. 4A). The specimen abundance (SA) shows the highest values in intermediate levels of the late Asbian, early Brigantian and early Serpukhovian, but with an overall upwardsdecreasing trend into younger intervals (Fig. 5A). In Ireland, although there are no upper Serpukhovian carbonates recorded in the succession, the total diversity (Fig. 4A) and the group diversity (Table 2; Supplementary Information Fig. S1B) follow similar Table 1 Statistical metrics evaluated in this study. Palaeoecological diversity includes both the D group and D total metrics. Metric Explanation Genus richness, GR number of genera for each substage Group diversity, D group The Margalef diversity index using the class divisions applied individually to the ten groups in section 2.2 Total diversity, D total The Margalef diversity index using the class divisions of genera in section 2.2 Specimen abundance, SA total number of identified specimens in each sample Volumetric abundance, %V Quantified in thin sections using the charts of Baccelle and Bosellini (1965) Taxonomic distinctness, Δ + Supragenerical divisions grouped by substage intervals following Clarke and Warwick (1998) P. C´ ozar et al. Earth-Science Reviews 246 (2023) 104596 4 Fig. 2. Schematic stratigraphic successions of the studied regions showing the predominant lithologies and environments. Coloured tie lines: green line, Asbian/ Brigantian boundary; red line, base of early Serpukhovian; blue line, base of late Serpukhovian. Abbreviations: e Asb =early Asbian; e Brig =early Brigantian; Krasno. =Krasnopolyanian; L =Lissylisheen Member (Slievenaglasha Formation); P-Z =Protvian-Zapaltyubian; Prot. Protvian; St. Steshevian; Taru =Tarusian; Vo =Voznesenkian; (see details of the stratigraphic sections and precise biostratigraphic information of Formation/Member/samples in Supplementary information and Appendix A). (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.) P. C´ ozar et al. Earth-Science Reviews 246 (2023) 104596 5 changes to those observed in northern England. The stratigraphical style during the Asbian is rather similar to that in northern England, with rhythmic limestones capped by emergent surfaces, whereas in Ireland a more continuous limestone succession is recorded for the early Brigantian and Serpukhovian (Fig. 2). Subtle differences are observed, such as lower dasycladal and calcifoliid D group values in the early Serpukhovian, which could be interpreted as local environmental differences (Fig. 4A). In Ireland, most of the algal associations recognised in northern England are also present, with the associations-1 A, 1B, 1C and 2 also occurring in the same stratigraphic order and nearly covering the same stratigraphic intervals (Fig. 4A). In Ireland the dasycladales association only occurs above the base of the Brigantian, although the presence of emergent surfaces at this level in Ireland implies a sedimentary gap, with the probable absence of the uppermost Asbian strata (C´ ozar et al., 2022c). The specimen abundance shows a decreasing trend from the early Brigantian, similar to that seen in the northern England data (Fig. 5A). Peaks in SA recognised in the late Asbian and base of the early Brigantian are synchronous between Ireland and northern England, whereas trends in both these regions differ during the Serpukhovian. 4.2. LPT (Sierra Morena and Adarouch) The successions in Sierra Morena and Adarouch are both mainly noncyclic in character, with that of Sierra Morena representing a continuous carbonate accumulation in the lower part of the succession (predominantly with microbially-mediated carbonates), whereas thick shale Fig. 3. Chronostratigraphical correlation of some regional substages (modified from Liu et al., 2023). British scale based on C´ ozar and Somerville (2021a, 2021b) and Hounslow et al. (2022) for the Mississippian, and Waters (2011) for the Pennsylvanian. Note that the base of the Holkerian and Asbian are modified (dotted lines). Scale from Belgium is based on Poty et al. (2006); East European Platform (EEP) from Russia is based on Alekseev et al. (2022); Donetz substages in Ukraine are based on Poletaev et al. (2013) and modification by Nemirovska (2017) for the Serpukhovian; the Chinese scale is based on Wang et al. (2019); and the American scale is based on the type series in the Mississippi River Valley in Illinois by Lane and Brenckle (2005). The dark shaded parts in the Britain and EEP scales are the selected regional substages division used in this study. Abbreviations: Arn Arnsbergian, Cha. Chadian, up. Upper. P. C´ ozar et al. Earth-Science Reviews 246 (2023) 104596 6 intervals are recorded between limestone intervals in Adarouch (suggesting four major deepening sequences and a final fifth shallowing sequence, Se1 to Se5 in Fig. 2). The late Asbian, mostly recorded in Sierra Morena, shows rather consistent values in total diversity (Fig. 4B), with the same algal components as observed in Laurussia, although with larger D group values for blue-green algae (Table 2; Supplementary Information Fig. S3A). The larger contribution from blue-green algal is a consequence of the common microbial facies in the inner platform (Fig. 2), a facies not present in the British/Irish inner platforms. The early Brigantian shows higher D total values (Fig. 4b) than in the late Asbian, and in contrast with Laurussia, the dominant groups of the late Asbian also extend into the early Brigantian (Supplementary Information Fig. S3A). The overall changes in D group in the Adarouch region are similar to those in Sierra Morena (Table 2; Supplementary Information Fig. S3B). Small differences are interpreted to be local characters, influenced by ecological or environmental conditions. In contrast with Laurussia, marked lows (D total <0.5) in the total diversity are scarcer (Fig. 4B), and generally D total values exceed 2 during the early Brigantian. The absence of these lows, as well as the absence of associations-2 and 3 during the early Brigantian, might be related to the absence of typical Yoredale-type cycles in the LPT, and thus the differing environmental and ecological conditions. The early Serpukhovian contains marked changes in D total (Fig. 4B), like those in Laurussia. These D total changes are more marked in Sierra Morena than in Adarouch, with slightly lower D total values in the upper part of this substage. The late Serpukhovian seems to show a recovery in the total diversity, reaching similarly high values (D total >1.5) to those during the early Brigantian, an effect mostly due to the dasycladales (Fig. 4B). During the Serpukhovian, basins analysed herein were tectonically active (e.g., C´ ozar et al., 2006, 2023b). The recognition of associations in the LPT is rather similar to that in Britain, and the association-1 A is recognised in the late Asbian of Adarouch (but is not observed in Sierra Morena), but the stratigraphic interval covered by each association differs from the Laurussia sections (Fig. 4). In term of abundances, in Sierra Morena, the late Asbian has consistently larger values (SA >10), whereas larger SA at Adarouch is more consistently located in the early Brigantian (Fig. 5B). In both cases, abundances generally decrease into the early Serpukhovian. The late Serpukhovian in Sierra Morena also contains large abundance values (Fig. 5B). Thus, the uppermost part of the succession in the LPT is different from the Laurussia Platform, with the peaks in Laurussian SA data during the late Asbian-Brigantian which are not observed in the LPT. This is perhaps the result of differing cyclic sedimentation patterns (Fig. 2). Both successions show a transition into deltaic facies in the upper part, at the top of the Protvian in Sierra Morena, and also at Adarouch, dated to the Voznesenkian (Fig. 2), where carbonates commonly contain quartz grains, representing more hostile facies for algae. Nevertheless, during the Bashkirian, representatives of the groups dasycladales, bluegreens, donezellids and rare aougaliids are recorded in these beds, belonging to the same genera/species seen in older Serpukhovian levels in the LPT. 4.3. GPT (Montagne Noire and Valdediezma) This region contains the thickest carbonate accumulation of all those in the study area (Fig. 2). Late Asbian strata in the Montagne Noire are poorly represented, with the lower parts of the substage commonly represented by siliciclastic turbidites (C´ ozar et al., 2017), or absent because of faulting in the Valdediezma Platform. This latter succession from northern Spain displays the oldest carbonates, dated to the upper part of the Mikhailovian Substage (=early Brigantian; Fig. 3) (BlancoFerrera et al., 2021). The short late Asbian interval in Montagne Noire contains smaller D total values than the Brigantian (Fig. 4C), as also observed in the LPT platforms. The negligible contributions of Koninckopora to the diversity are a remarkable and unique feature, which characterises the GPT (Table 2; Supplementary information Fig. S4). The early Brigantian shows a similar scale of fluctuations in total diversity as in other regions (Fig. 4), and the most important groups are similar to those in the Asbian elsewhere in Laurussia and LPT (Table 2). D total values >1.5 in the early Serpukhovian in the GPT are in marked contrast with the LPT, because these values exceed those during the early Brigantian (Fig. 4B, C). This represents a somewhat similar relationship to that observed in northern England. The incoming of dasycladales is the main group contributing to the higher D group values in the GPT (Supplementary Information Fig. S4). The late Serpukhovian contains high D total values for the lower part (equivalent to the Protvian), and a marked decrease for the upper part (equivalent to the Zapaltyubian) at Valdediezma (Fig. 4C). Similar variation in D total in the upper part of the Montagne Noire succession is not observed, because the interval corresponding to the Zapaltyubian is composed of siliciclastic deltaic deposits and algae were not recorded (C´ ozar et al., 2019). The GPT contains the most unusual algal associations, and only associations 1B, 3 and 4 are found (Fig. 4C), with no similarities between sections, or with the other regions. In terms of algal abundance, smaller values occur during the Table 2 Dgroup and main contributor taxa. late Asbian early Brigantian early Serpukhovian late Serpukhovian Northern England high (Koninckopora, donezellids, Ungdarella, aoujgaliales, moravamminids) moderate (dasycladales, early peaks of calcifoliids) moderate (calcifoliids, aoujgaliales, dasycladales) very low Ireland high (donezellids, Koninckopora, Ungdarella, aoujgaliales, moravamminids) moderate (dasycladales, aoujgaliales) moderate to low, fluctuating (aoujgaliales, calcifoliids) Absent Sierra Morena high (donezellids, Koninckopora, calcifoliids, moravaminids, bluegreens high (donezellids, Koninckopora, aoujgaliales, calcifoliids, moravamminids, blue-green moderate (aoujgaliales, calcifoliids, blue-greens, rhodophytes) moderate (blue-greens, calcifoliids, aoujgaliales, dasycladales) Adarouch poor (Koninckopora, aoujgaliales) high (cyanobacteria, Ungdarella, aoujgaliales, donezellids) moderate (calcifoliids, aoujgaliales, blue-greens) moderate to low (calcifoliids, aoujgaliales, blue-greens) Montagne Noire moderate (donezellids, calcifoliids, Ungdarella) moderate to low (calcifoliids, donezellids, aoujgaliales) high (calcifoliids, donezellids, bluegreens, dasycladales, bryopsidales) moderate (calcifoliids, dasycladales, blue-greens, donezellids) Valdediezma absent moderate (calcifoliids, donezellids, bluegreens moderate to high (calcifoliids, bluegreens, dasycladales, donezellids) moderate (calcifoliids, donezellids,blue-greens, dasycladales) Tindouf moderate (Koninckopora, aoujgaliales) moderate (koninckopora, aoujgaliales) moderate (Aoujgaliales, Koninckopora, dasycladales) low (Koninckopra, donezellids, dasycladales) P. C´ ozar et al. Earth-Science Reviews 246 (2023) 104596 7 Brigantian, increasing towards the top of the early Serpukhovian, then generally decreasing to the top of the GPT successions (Fig. 5C). The succession in the Valdediezma Platform contains well-developed early Bashkirian carbonates (Fig. 2), interrupted by an overlying unconformity, onlapped by Moscovian carbonates. In the early Bashkirian units, total diversity is <1.0, with mostly calcifoliids and donezellids occurring. These are noteworthy for the occurrence of Donezella (Fig. 6H), a typical Pennsylvanian taxon, which is the only renovation within the GPT assemblages. 4.4. Gondwana (Tindouf) The succession in the Tindouf Basin is distinctly cyclic from the early Brigantian onwards (Fig. 2), although complete cyclothems veneered by coal were not developed (C´ ozar et al., 2014). This platform shows large total diversity (generally D total >1.5) during the late Asbian and early Brigantian (Fig. 4D). The early Serpukhovian has more D total values, largely >2, with often total diversity <1 for the late Serpukhovian (Fig. 4D). The overall changes in D total are similar to those observed in Laurussia, with both regions showing an overall decreasing trend into the late Serpukhovian. In terms of group diversity, Koninckopora largely makes a major contribution throughout the succession, much like that shown by aoujgaliales (Table 2; Supplementary Information Fig. S5). Negligible contributions are from blue-green algae, whereas the dasycladales form a smaller contribution to the diversity, which is particularly important during the early Serpukhovian. These features are also observed in the Laurussian platforms. The succession in Tindouf contains five of the six algal associations, with only the association-1C not recognised (Fig. 4D). The timings of the Fig. 4. Margalef diversity index for total algal diversity (D total ). Abbreviation: e. early. Taru. Tarusian, Stesh. Steshevian, Prot. Provian, Zapalt. Zapaltyubian, Voz. Voznesenkian, Krasno. Krasnopolyanian. LPT =Laurussian Palaeotethys terranes, GPT =Gondwana Palaeotethys terranes. Y-scale =ordered stratigraphic position of samples. Database included in Appendix A. P. C´ ozar et al. Earth-Science Reviews 246 (2023) 104596 8 appearance of associations shows a marked delay compared to the British/Irish data (Fig. 4). The interval equivalent to the Zapaltyubian and Voznesenkian in the Tindouf succession are composed of sandy supratidal sabkhas (Fig. 2), with rare occurrence of algae. However, in the upper part of the carbonate succession (assigned to the Krasnopolyanian Substage), there is a recovery in the total diversity and abundance of algae (Figs. 4D, 5D). The main components in the Krasnopolyanian age beds are aoujgaliids (mostly Aoujgalia), Ungdarella, calcifoliids (Calcifolium and Falsocalcifolium) and donezellids (Praedonezella). These genera are the same as those from the Serpukhovian in Tindouf, and thus there are no new typically Pennsylvanian taxa. 5. Volumetric contribution (%V) of calcareous algae to the carbonate factory The late Asbian is the interval with the largest contribution made by calcareous algae, forming 40% to 70% of the total content in beds (including allochems, matrix and cement) (Fig. 7). The fluctuations in % V values in the Laurussia Platform successions seem to be controlled by glacioeustatic cycles (Fig. 7A), which are rather similar to the foraminiferal trends defined between emergent surfaces in Britain and Ireland (C´ ozar et al., 2022c). Surprisingly, at Sierra Morena, virtually free of emergent surfaces for this interval and with no recognisable cyclicity, similarly-timed peaks of similar algal composition and cycles are also observed (Fig. 7A, B). However, other lithologically non-cyclic Fig. 5. Specimen abundance (SA). Abbreviation: e. early. Taru. Tarusian, Stesh. Steshevian, Pro. Protvian, Zapalt. Zapaltyubian, Voz. Voznesenkian. See Fig. 3 for other details. P. C´ ozar et al. Earth-Science Reviews 246 (2023) 104596 9 Fig. 6. Algal associations (scale bar =1 mm). A. Kamaenella (white arrows)-Koninckopora (k) association, sample 1274, early Asbian, Tankardstown Borehole, Ireland; Kamaenella has been arrowed in a small part of the picture, but the entire small rounded sections in the matrix correspond to this genus. B. Kamaenella (white arrows)-Koninckopora (k) association, sample TQ-205, late Asbian, Trowbarrow Quarry, northwest England; only a few Kamaenella have been arrowed. C. Kamaenella (white arrows)-Ungdarella (u)-Koninckopora (k) association, sample TQ-75, late Asbian, Trowbarrow Quarry, northwest England; only a few Kamaenella have been arrowed. D. Ungdarella (u)-Kamaenella (k) association, sample TQ-139, late Asbian, Trowbarrow Quarry, northwest England. E. Dasycladales association (w Windsoporella, c Coelosporella, n Nanopora), sample PC918, early Brigantian, Clogrenan Quarry, Ireland. F. Falsocalcifolium association, late Serpukhovian, La Serre, Montagne Noire (France). G. Calcifolium association, late Serpukhovian, Jitu l’Escarandi, Cantabrian Mountains (Spain). H. Facies with Donezella (white arrows), early Bashkirian, Jitu l’Escarandi, Cantabrian Mountains (Spain); only a few Donezella have been arrowed, but most rounded forms correspond to this taxon, which is rather recrystallized. P. C´ ozar et al. Earth-Science Reviews 246 (2023) 104596 16 (Fig. 10A), coinciding with a widespread hiatus, but without clearly defined cyclic sequences. Following this event is an overall increase in taxonomic diversity of some fossil groups, and marked ecosystem changes in both foraminiferal and algal diversity. These changes and reduction of the diversity are recorded in Britain/Ireland in horizons in the uppermost part of this interval (Fig. 10A, B). However, as discussed above, a tectonic control on this hiatus seems to be a better explanation. (Event 2): At the onset of cyclic-rhythmic sequences at the base of the late Asbian, which also coincides with ecosystem changes (Fig. 10A). This event is not free of controversy, since in Britain at least, apart from possible cumulative autogenic sedimentation processes, local/regional tectonics and eustasy have all been suggested as originators of these rhythms/cycles (see revision in Horbury, 1987). The inference has been, that in the sections in NW England, the lower five of these rhythms were considered tectonically-driven cycles by Horbury (1987, 1989). There is certainly a larger influence of local tectonics in these older cycles which are not veneered by terrestrial deposits in some parts of Ireland and Wales (C´ ozar et al., 2022c). The consistent foraminiferal diversity changes attributed to glacioeustasy also confirm the influence of this factor from the base of the late Asbian. Assuming a tectonic control on the lower five cycles, the occurrence of the overlying cycles of entirely glacioeustatic-origin would then be situated in the Cf6γ2 British foraminiferal subzone (Fig. 3), a level which is difficult to identify and calibrate outside of Britain and Ireland. (Event 3): At the base of rather fluctuating biotic-responses close to the base of the early Serpukhovian, which coincides with the Main Eustatic Signal (MES) of Fielding and Frank (2015), also recognised in some North American basins (Ahern and Fielding, 2019, 2021); which post-dates the Marine Ecosystem Resurgence (MER) (Fig. 10A, B). (Event 4): At the onset of the main taxonomic turnover of the biota in the latest Serpukhovian, which coincides with sustained changes in δ 18 O values of brachiopods, and the ending of the C1 glacial phase of Fielding et al. (2008a, 2008b) (Fig. 10A, B). Owing to its likely tectonic control, event 1 is not of the same worldwide significance (C´ ozar et al., 2023a). Event 2, the onset of the cyclic sedimentation as a result of glacioeustasy at the base of the late Asbian, needs to be further investigated to solve problems in biostratigraphy, and precisely constrain the synchronicity of the onset of glacioeustatic cycles, or onset of high-amplitude sea-level falls. The possible influence of regional tectonics in some basins needs to be better understood. Event 3 seems to be synchronous in terms of biota changes, and coincides with the MES, and its post-dating of the MER. Event 4 coincides with the end of glacial phase C1, but also with major changes in ocean water circulation due to the closure of the Rheic-Palaeotethys gateway. This stimulated stronger weathering of silicates leading to water chemistry changes, and narrower shallow-marine habitats (Sandberg, 1983; Hardie, 1996; Morse et al., 1997; Davydov and C´ ozar, 2019). Therefore, it is questionable whether a ‘sustained glaciation’ can be applied in the interval of the LPIA until the Serpukhovian. This is because far-field proxies are inconsistent with each other (such as cyclicity), and isotopic signatures progressively change during the Mississippian, without clearly defined cooling events. In terms of biota, an important change occurred in ecosystems during the early Serpukhovian, whereas taxonomic turnovers started in the uppermost Serpukhovian. In consequence, to establish a point in the middle of a continuum of isotopic enrichment, or thresholding what is a significant sea level fall (i.e. >25 m) does not firmly establish the onset of sustained icehouse conditions. The time when progressive cooling affected the biota in equatorialtropical regions is a more pragmatic level for the definition of when global icehouse conditions were first sustained in the LPIA. In contrast, if commonly inferred times of icehouse initiation are used, such as the base of the middle Vis´ ean (339 Ma approx.) and late Asbian (334 Ma approx.), it is unclear why the main ecosystem change occurred during the early Serpukhovian (330.34 Ma to 327 Ma approx.), and taxonomic changes in the most sensitive biota occurred from the uppermost Serpukhovian to the early Bashkirian (around 323.4 Ma). This is some 4 to 17 Myr later (depending on the selected levels) than the assumed onset of icehouse conditions in the middle Vis´ ean or late Asbian. If greenhouse to icehouse transitions typically occurred over durations of 1–1000 kyrs (Saupe et al., 2020), then either the duration of such brief transitions is incorrect for the Mississippian, or the onset of sustained icehouse conditions should be restricted to the Serpukhovian. 11. Conclusions Study of the calcareous algae from the late Vis´ ean to Serpukhovian interval in low-palaeolatitude basins from the western margin of the Palaeotethys show diverse responses in terms of palaeoecological diversity, richness, abundance, genus diversity and taxonomic distinctness. Conclusions inferred from these changes are: - Algal associations are more similar in cratonic areas, although many diversity peaks correspond to local ecological/environmental factors. - The Laurussian Palaeotethyan terranes contain the most unusual algal abundances and are the most affected by active tectonics and palaeogeographic reorganization. - Algae in palaeo-equatorial regions show more dramatic changes than in the palaeo-tropical regions, which are possibly more rapidly affected by the reduction of continental shelves. - Genus richness shows an inflexion in the curve of originations/extinctions in the late Vis´ ean-early Serpukhovian interval. - Taxonomic distinctness decreases significantly from the uppermost Serpukhovian. - The variety of algal diversity responses suggests that it is not appropriate to extrapolate particular diversity responses from a single region to global conditions, and many factors likely influenced the regional assemblages. - The main taxonomic turnover of most fossil groups started from the uppermost Serpukhovian, likely linked to a significant phase of cooling based on δ 18 O from brachiopods. This coincides with final stages of the glacial phase C1 defined in Australian basins, but most palaeo-equatorial ecological changes occurred from the early Serpukhovian, coinciding with the Main Eustatic Signal seen in British and North American basins. - Foraminifers and algae show rather similar ecological diversity for the Vis´ ean and Serpukhovian in Britain, with a single coincident inflexion point in their trends in the early Serpukhovian. Far-field proxies defining the onset of icehouse period in the LPIA are inconsistent in timing, and are located in distinct stratigraphical levels far from the main biota changes. This questions the usefulness of the previously inferred onset of sustained icehouse glaciation for the palaeoequatorial Vis´ ean, and a better level to define this major event was when low-palaeolatitude biota suffered the effects of water cooling during the Serpukhovian. CRediT authorship contribution statement Pedro C´ ozar: Conceptualization, Resources, Investigation, Methodology, Formal analysis, Writing – original draft. Ian D. Somerville: Investigation, Resources, Writing – original draft. Mark W. Hounslow: Resources, Writing – review & editing. J. Ricardo Mateos-Carralafuente: Writing – review & editing. Ismael Coronado: Investigation, Methodology, Writing – original draft. Declaration of Competing Interest None. P. C´ ozar et al. Earth-Science Reviews 246 (2023) 104596 17 Data availability All data used are presented in the Supplementary files. Acknowledgements We thank the editor (C. R. Fielding) and two reviewers for their many helpful comments and suggestions which have significantly improved the paper. Appendix A. Supplementary data Supplementary data to this article can be found online at https://doi. org/10.1016/j.earscirev.2023.104596. References Adams, A.E., 1984. 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