Biodiversity of ecosystems in an arid setting: The late Albian plant communities and associated biota from eastern Iberia

RESEARCH ARTICLE Biodiversity of ecosystems in an arid setting: The late Albian plant communities and associated biota from eastern Iberia Eduardo Barro´ nID1*, Daniel Peyrot2, Carlos A. Bueno-Cebollada1☯, Jiřı´ Kvaček3, Sergio A´ lvarez-ParraID4,5, Yul Altolaguirre6, Nieves Mele´ ndez7☯ 1 Museo Geominero, Centro Nacional Instituto Geolo´ gico y Minero de España CN-IGME CSIC, Madrid, Spain, 2 School of Earth and Environment, Centre for Energy Geoscience, University of Western Australia, Crawley, Western Australia, Australia, 3 National Museum Prague, Praha, Czechia, 4 Departament de Dinàmica de la Terra i de l’Oceà, Facultat de Ciències de la Terra, Universitat de Barcelona, Barcelona, Spain, 5 Institut de Recerca de la Biodiversitat (IRBio), Universitat de Barcelona, Barcelona, Spain, 6 OCEEH Research Center ‘The role of cultura in early expansions of humans’, Heidelberg Academy of Sciences, Senckenberg Research Institute, Frankfurt am Main, Germany, 7 Deparment of Geodina´ mica, Estratigrafı´a y Paleontologı´a, Facultad de Ciencias Geolo´gicas, Universidad Complutense de Madrid, Madrid, Spain ☯ These authors contributed equally to this work. * e.barron@igme.es Abstract Deserts are stressful environments where the living beings must acquire different strategies to survive due to the water stress conditions. From the late Albian to the early Cenomanian, the northern and eastern parts of Iberia were the location of the desert system represented by deposits assigned to the Utrillas Group, which bear abundant amber with numerous

bioinclusions, including diverse arthropods and vertebrate remains. In the Maestrazgo Basin (E Spain), the late Albian to early Cenomanian sedimentary succession represents the most distal part of the desert system (fore-erg) that was characterised by an alternation of aeolian and shallow marine sedimentary environments in the proximity of the Western Tethys palaeo-coast, with rare to frequent dinoflagellate cysts. The terrestrial ecosystems from this area were biodiverse, and comprised plant communities whose fossils are associ- ated with sedimentological indicators of aridity. The palynoflora dominated by wind-trans- ported conifer pollen is interpreted to reflect various types of xerophytic woodlands from the hinterlands and the coastal settings. Therefore, fern and angiosperm communities abun- dantly grew in wet interdunes and coastal wetlands (temporary to semi-permanent freshwa- ter/salt marshes and water bodies). In addition, the occurrence of low-diversity megafloral assemblages reflects the existence of coastal salt-influenced settings. The palaeobotanical study carried out in this paper which is an integrative work on palynology and palaeobotany, does not only allow the reconstruction of the vegetation that developed in the mid-Creta- ceous fore-erg from the eastern Iberia, in addition, provides new biostratigraphic and palaeogeographic data considering the context of angiosperm radiation as well as the biota inferred in the amber-bearing outcrops of San Just, Arroyo de la Pascueta and La Hoya

(within Cortes de Arenoso succesion). Importantly, the studied assemblages include Afro- pollis, Dichastopollenites, Cretacaeiporites together with pollen produced by Ephedraceae PLOS ONE PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 1 / 51 a1111111111 a1111111111 a1111111111 a1111111111 a1111111111 OPEN ACCESS Citation: Barro´n E, Peyrot D, Bueno-Cebollada CA, Kvaček J, A´lvarez-Parra S, Altolaguirre Y, et al. (2023) Biodiversity of ecosystems in an arid setting: The late Albian plant communities and associated biota from eastern Iberia. PLoS ONE 18(3): e0282178. https://doi.org/10.1371/journal. pone.0282178 Editor: Enrique Peñalver, Geological and Mining Institute of Spain, SPAIN Received: July 29, 2022 Accepted: February 8, 2023 Published: March 2, 2023 Peer Review History: PLOS recognizes the benefits of transparency in the peer review process; therefore, we enable the publication of all of the content of peer review and author responses alongside final, published articles. The editorial history of this article is available here: https://doi.org/10.1371/journal.pone.0282178 Copyright: © 2023 Barro´n et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Data Availability Statement: All relevant data are within the paper and its Supporting Information files.

(known for its tolerance to arid conditions). The presence of these pollen grains, typical for northern Gondwana, associates the Iberian ecosystems with those characterising the men- tioned region. Introduction The Albian vegetation of western Europe has been studied in several works which described micro-, meso- and macroremains [1–7], and highlighting the prevalence of ferns and gymno- sperms–mainly conifers (Araucariaceae-Podocarpaceae-Pinaceae-Cupressaceae-Cheirolepi- diaceae)–, and the increasing representation of angiosperms. The distinctiveness of the western European flora has been recognised for a long time and several biogeographic prov- inces have been named on the basis of the nature of the fossil assemblages (i.e., palynological and megaremains [8, 9]). The increased attention to the time-interval is justified by the rising of the dominance of the flowering plants in many ecosystems [10–13] including arid environ- ments [14]. During the later part of the Early Cretaceous, SW Europe has been described as one of the probable migratory routes of angiosperms originating from a northern Gondwanan centre of radiation [15]. The successive northward migrations of angiosperms, well-character- ised by both pollen and megaremain records, suggest broadly similar climate conditions in low to mid-latitude settings (i.e., low latitudinal climatic gradient) or selective migration of taxa able to tolerate a wide range of climatic conditions [16]. During most of the Albian, Iberia was part of a climatic zone referred to as the Northern Hot Arid (NHA) belt [17], where a desert

system (erg) developed over an extensive area [18]. Arid environments, although less represented in fossil record, existed during the Mesozoic. Semi-arid environments are, e.g., reconstructed for the Early Triassic of northeastern Spain and the Upper Jurassic Morrison Formation based on presence of aeolian sandstones [19, 20]. Arid environments were inferred for Early Cretaceous floras of Quedlinburg (Germany) and Las Hoyas (Central Spain) based on occurrence of dwarf lycopod Nathorstiana and fern Weichselia [21, 22]. Reconstructions of Cretaceous continental ecosystems of Japan based on stable oxygen and carbon isotopes also suggest semi-arid conditions [23]. In addition, aeolian sandtones provide direct evidence of a desert environment during the mid-Cretaceous in mid- to low latitude Asia [24]. In Iberia, the eastern coastal margin consisted of a complex set of depositional environ- ments comprising multifaceted vegetation types including conifer forests [14, 25]. Palaeobota- nical studies of ancient desert successions (see i.e. [26, 27]) report plant remains usually not preserved in sand-sized lithologies, which in turn constitute the bulk of their deposits. Assem- blages belonging to this type of setting usually include remains that have experienced transport over varying, and sometimes considerable, distances to the site of burial. Considering this limi- tation and exploiting the characteristics of sets of fossils with distinct taphonomic histories (i.e., megaremains with limited transport vs. miospores disseminated over wider distances),

the present work focuses on the reconstruction of the mid-Cretaceous plant communities that developed in the distal part of the desert system (or fore-erg) in the Maestrazgo Basin (Eastern Iberian Ranges, E Spain), considering specially the conifer-dominated ones. While the previ- ous studies carried out in Albian deposits from the Maestrazgo Basin considered palaeobotani- cal and palynological data separately [28–37], this study represents the first attempt to integrate both data sets into a unified framework, aiming at the characterisation of the arid palaeoenvironmental and their link with the occurrence of amber. The Cretaceous amber-bearing outcrops from the Maestrazgo Basin have been known for a long time [38] and were traditionally associated with the Escucha Formation [39], an Aptian– PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 2 / 51 Funding: This study is a contribution to the project CRE CGL2017-84419 AEI/FEDER, UE from the Ministerio de Ciencia, Innovacio´n y Universidades (Spain) and the “Severo Ochoa” extraordinary grants for excellence IGME-CSIC (AECEX2021). The coauthor S.A´.-P. thanks the support from the Secretaria d’Universitats i Recerca de la Generalitat de Catalonia (Spain) and the European Social Fund (2021FI_B2 0003). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing interests: The authors have declared that no competing interests exist.

early Albian unit consisting of coal-bearing deposits [40] which sometimes also present amber [41]. Although more than 30 amber outcrops have been reported in the basin, the sites related to the desert system revealing the most diverse sets of bioinclusions and plant remains [29, 37, 42–44] are the successions of Arroyo de la Pascueta, San Just (both located in Teruel Province) and Cortes de Arenoso (Castello´n Province) (Fig 1). The present study is the first to provide a comprehensive treatment of the palaeobotanical and palynological content of the three afore- mentioned mid-Cretaceous sites. This paper aims to: (i) document the micro- and macrofloral assemblages together with the entomological content recovered from highly significant amber- bearing outcrops; (ii) reassess the age of the successions based on the presence of key biostrati- graphic markers; (iii) reconstruct the corresponding ecosystems with a special focus on the plant communities and insects which thrived on them; and (iv) compare the vegetation with coeval floras from presumably wetter settings. Material and methods Geological setting The study area is located in the Maestrazgo Basin, on the eastern part of a palaeogeographic domain referred to as the Iberian Basin Rift System (IBRS), Spain (Fig 1A). The sedimentary Fig 1. Location of the Maestrazgo Basin and its seven sub-basins (A) based on Aurell et al. [45]. The studied locations are indicated by coloured stars (San Just,

SJ; Arroyo de la Pascueta, AP; Cortes de Arenoso, CA). (B) Detail of the geological map with the location of San Just (SJ) section in the Las Parras Sub-basin. Based on data from the geological map of Spain Magna 1:50000 [46]. (C) Detail of the geological map showing the location of Arroyo de la Pascueta (AP) and Cortes de Arenoso (CA) sections. Based on data from the geological map of Spain Magna 1:50000 [47]. https://doi.org/10.1371/journal.pone.0282178.g001 PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 3 / 51

succession was deposited during the transition from the Late Jurassic–Early Cretaceous syn- rift stage to the Late Cretaceous post-rift stage, associated with the gradual opening of the Western Tethys and the North Atlantic Ocean [45, 48, 49]. The Maestrazgo Basin forms a NW-SE-oriented depositional trough (Fig 1A), where the sedimentary succession generally becomes thicker towards the SE (Tethyan margin). During the Early Cretaceous the syn-rift sedimentary succession was dominated by thick, continental to coastal siliciclastic-dominated deposits (previously referred to as ‘Weald facies’) and marine carbonates (‘Urgonian facies’) [50–55] (Fig 2). Towards the end of the syn-rift stage (uppermost Aptian to early Albian), the sedimentary succession became more homogeneous across the basin, and coal-bearing strata attributed to the Escucha Formation were deposited [56–59]. The Escucha Formation has been interpreted as lagoons and marshes [59, 60]. The Escucha Formation is overlain by strata forming the Utrillas Group, a highly diachro- nous unit widespread along the IBRS, which has been traditionally interpreted as the transition into the post-rift stage [57, 59, 61]. In the Maestrazgo Basin, the Utrillas Group spanned the middle Albian–earliest Cenomanian [59]. In the Maestrazgo Basin, this Group includes facies originally interpreted as tidally-influenced fluvial deposits [57, 62] and later reinterpreted as part of an erg system (aeolian dune desert), covering more than 16,000 km2 on the eastern side of the Iberian Peninsula [18, 59].

According to [18, 59], this erg system (Fig 2B) displayed a threefold zonation developed along a NW–SE direction characterised by: (i) a back-erg area close to the main source area of siliciclastics (Iberian Massif), dominated by coarser-grained wadi sediments and minor aeolian accumulations; (ii) a central-erg area corresponding to the main locus of aeolian dune accu- mulation; and (iii) a fore-erg area characterised by the interplay between aeolian dunes and shallow marine deposits from the Western Tethys, giving rise to a complex interaction of tid- ally-reworked aeolian dune and subtidal facies associations. In the Maestrazgo Basin, the Utrillas Group is overlain by the Mosqueruela Formation [50, 57, 59, 63] (Fig 2B), a marine carbonate unit, that represents the onset of carbonate sedimenta- tion (Southern Iberian Ramp), which will prevail during most of the Late Cretaceous [48, 64]. The Maestrazgo Basin is divided into several sub-basins (Fig 1) that include: the Galve, Oliete, Las Parras, Morella, El Perello´, La Salzedella, and Peñagolosa sub-basins [49]. In this paper, three stratigraphic sections have been measured in three outcrops in the Maestrazgo Basin, namely: San Just (according to [31]: “left slope of San Just”), Arroyo de la Pascueta and Cortes de Arenoso (according to [40]: “La Hoya outcrop”) sections. The San Just section is located in the Las Parras Sub-basin, about 70 km to the NNW from the other studied sections which crop out in the NNW sector of the Penyagolosa Sub-basin (Fig 1). The studied

sedimentary succession is located in the distal part of the mid-Cretaceous desert system, which is characterised by a fore-erg setting where aeolian strata interacted with shallow marine deposits in the coastal fringe of the Western Tethys. San Just section (SJ). The 22 m-thick section is located on a roadside (N-420 road), approximately 3 km to the south of Utrillas village (Teruel Province) and overlain by the marine carbonates of the Mosqueruela Formation, dated as Cenomanian [46]. The succession can be divided into a lower part mainly consisting of dark siltstones and mudstones inter- spersed with cm- to dm-thick fine-grained sandstone layers with wavy to flaser heterolithic laminations (Fig 3), and an upper part dominated by fine-grained, cross-bedded, sandstones with thin (dm-thick) interbedded mudstones, which may include water deformation struc- tures and slumps. The amber-bearing bed is located in the lower part of the section and is over- lain by (large-scale) planar cross-bedded fine-grained sandstones exhibiting sharp basal contacts. Kindney-shaped amber pieces together with “stalactitic aerial” amber pieces, the lat- ter with bioinclusions, are abundant in this bed [43] (Fig 4A and 4B). PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 4 / 51

Cortes de Arenoso section (CA). This 172 m-thick section is cropping out in a ravine in the NNW of the Cortes de Arenoso locality (Castello´n Province). The succession, sharply over- lain by marine carbonates attributed to the Mosqueruela Formation [47, 65], is dominated by fine-grained sandstones displaying large-scale planar cross-bedded sets, which laterally shift into dark mudstones and siltstones containing plant megaremains, especially abundant in the first 10–30 m of the logged section (Fig 4). The section includes heterolithic siltstone/mud- stone and sansdtone deposits displaying subaqueous tidal sedimentary structures such as dou- ble mud drapes, lenticular, flaser and wavy laminations and cross-bedded sets depicting current bipolarity (herringbone-like cross-beddings). The amber-bearing level corresponds to the La Hoya amber outcrop (not to confuse it with the Barremian compression outcrop of Las Hoyas, also in Spain). Concretely, this level is located in the uppermost mudstone interval of the section and correlates with the amber-bearing stratum described in Arroyo de la Pascueta [40, 44]. Fig 2. Synthetic stratigraphic log of the Maestrazgo Basin from the Triassic to the Upper Cretaceous. (A) The red arrow shows the location of the Utrillas Group in grey colour. Based on data from [52, 55]. (B) Detail of the lithostratigraphy of the studied succession, where the Utrillas Group Erg System overlies the coal-bearing deposits of the Escucha Formation with a regional unconformity surface (RUS), which marks the onset of the Erg System. The Utrillas Group

succession is, in turn, overlain by marine carbonate deposits of Mosqueruela Formation. Based on data from [59]. https://doi.org/10.1371/journal.pone.0282178.g002 PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 5 / 51

Arroyo de la Pascueta section (AP). The 182 m-thick section consists of an alternation of fine-grained homometric sandstone bodies with large-scale planar cross-bedding (Fig 6). The sandstone laterally shifts into dark siltstone to mudstone and heterolithic deposits with mud drapes and wavy to flaser laminations, showing similar characteristics to the heterolithic silt- stone/mudstone described in the Cortes de Arenoso section (Fig 5). Despite plant remains being common and evenly distributed throughout the succession, only the lower part of the section (interval between 20 to 27 m in Fig 6) was sampled and analysed in previous Fig 3. Stratigraphic log of the upper Albian San Just outcrop (SJ), showing the distribution of selected biostratigraphically relevant palynomorphs. The San Just amber outcrop is at the top of the level 3. Red dots: location of the collected samples. Black dots: stratigraphic distribution of the selected taxa. Ø: barren sample. https://doi.org/10.1371/journal.pone.0282178.g003 PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 6 / 51

palaeobotanical studies [29, 30]. Amber fragments are abundant [40, 44] and recorded in a mudstone bed of the upper part of the succession, which is sharply overlain by the marine car- bonates of the Mosqueruela Formation [47]. Fig 4. Amber from the late Albian San Just outcrop (Utrillas, Teruel) and its bioinclusions. (A) Big kidney-shaped amber mass of 640 g. (B) Unearthed amber pieces. (C) Undetermined Lagonomegopidae? spider (SJNB2012-12-07). (D) Holotype of Actenobius magneoculus (Coleoptera, Ptinidae) (MAP-7727). (E) Holotype of Cretevania alcalai (Hymenoptera, Evaniidae) (CPT-960). (F) Avian dinosaur feather barb close to an undetermined Platygastroidea wasp (CPT-4078 and CPT-4079, respectively). Scale bars C–F, 0.5 mm. Images A and B are provided by Enrique Peñalver. Acronyms SJNB, CPT and MAP correspond to amber pieces housed at the Museo Aragone´s de Paleontologı´a (Fundacio´n Conjunto Paleontolo´gico de Teruel-Dino´polis) in Teruel, Spain. https://doi.org/10.1371/journal.pone.0282178.g004 PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 7 / 51

Palynology Ninety-two samples have been collected in the three cropping out successions described above (Fig 1). Although sampling primarily focused on amber-bearing strata, all suitable levels of the sections were carefully sampled (Figs 3–5) in order to obtain a more accurate and continuous biostratigraphic framework. The three logged sections were drawn using the software SedLog [66] in combination with Adobe Illustrator (www.adobe.com). A total of 15 rock samples from the San Just, 36 from the Cortes de Arenoso and 41 from the Arroyo de la Pascueta sections were prepared for palynological analysis. Palynological resi- dues were processed using acid digestion with HCl and HF at high temperatures [67, 68]. When required, a short oxidation with HNO3 (“nitric wash”) was performed in some residues. The residues were then concentrated by sieving through 500, 250 and 10 μm sieves, mounted Fig 5. Stratigraphic log of the upper Albian/lower Cenomanian Cortes de Arenoso section (CA), showing the distribution of selected biostratigraphically relevant miospores (see the legend in Fig 3). The La Hoya amber outcrop is at the top of the section (level 35). Red dots: location of the collected samples. Black dots: stratigraphic distribution of the selected taxa. Ø: barren samples. https://doi.org/10.1371/journal.pone.0282178.g005 PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 8 / 51

in glycerin jelly on strew slides. In order to include rare taxa with potential biostratigraphic value, between 500 and 1000 palynomorphs were identified per sample (S1 Appendix). Micro- scopic analysis of the palynological slides was performed with an Olympus BX51 microscope, incorporating a ColorView IIIu camera using a 100X oil immersion objective. Pollen diagrams were constructed by using Tilia/TGView 2.0.2 softwares [69, 70]. All studied slides are provi- sionally stored in the museum of the Geological Survey of Spain (CN IGME-CSIC, Madrid). A list of the identified taxa along with their botanical affinity and occurrences in each section is given in Table 1 and S1 and S2 Appendices. Fig 6. Stratigraphic log of the upper Albian Arroyo de la Pascueta section (AP), showing the distribution of selected biostratigraphically relevant miospores (see the legend in Fig 3). The Arroyo de la Pascueta amber outcrop is at the top of the section (level 34). Red dots: location of the collected samples. Black dots: stratigraphic distribution of the selected taxa. Ø: barren samples. https://doi.org/10.1371/journal.pone.0282178.g006 PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 9 / 51

Table 1. List ordered alphabetically of the palaeobotanical taxa considering their botanical affinities, and possible preferred environment which are based in biblio- graphic references for aquatic palynomorphs [71, 72], ferns and allied [73–76], gymnosperms [12, 24, 74, 77–91] and angiosperms [12, 92–99]. Taxon Section Known or probable parent affinity Possible preferred environment Abietinaepollenites sp. SJ Gymnospermophyta; uncertain affinity Canopy/xerophytic (upland, well drained) Acritosporites cf. kyrtomus CA Monilophyta; uncertain affinities (Lygodiaceae??) Ground cover/hygrophytic? Acritosporites sp. SJ Aequitriradites spinulosus CA Hepaticophyta Ground cover/xerophytic? Aequitriradites sp. AP Afropollis jardinus SJ, CA, AP Gymnospermophyta; uncertain affinity Canopy?, shrub?/xerophytic (lowland?) Antulsporites clavus SJ Bryophyta; Sphagnales; Sphagnaceae Ground cover/hygrophytic (marshes, peat bog?) cf. Antulsporites sp. AP Bryophyta?; Sphagnales? Ground cover/hygrophytic (marshes, peat bog?) Alisporites bilateralis CA, AP Gymnospermophyta; uncertain affinity (Corystospermales?, Podocarpaceae?) Canopy/xerophytic (coastal?, upland?) Alisporites grandis CA, AP Alisporites spp. CA Appendicisporites bifurcatus AP Monilophyta; Schizaeales; Anemiaceae Ground cover, understory, epiphytes/ hygrophytic (lowland, marshes, wet environments) Appendicisporites bilateralis SJ, CA, AP Appendicisporites cf. concentricus CA, AP Appendicisporites cf. crenimurus CA Appendicisporites cristatus CA Appendicisporites erdtmanii CA, AP

Appendicisporites fucosus CA, AP Appendicisporites cf. fucosus SJ Appendicisporites cf. jansonii CA, AP Appendicisporites potomacensis CA Appendicisporites cf. potomacensis AP Appendicisporites problematicus AP Appendicisporites cf. problematicus CA, AP Appendicisporites robustus CA Appendicisporites tricornitatus CA, AP Appendicisporites cf. tricornitatus CA Appendicisporites spp. SJ, CA, AP ?Apteodinium sp. CA Dinoflagellata: Gonyaulacaceae Marine Araucariacites australis SJ, CA, AP Gymnospermophyta; Coniferales; Araucariaceae Canopy/xerophytic (coastal, upland?) Artiopollis praecox CA, AP Antophyta; non-magnoliid dicotyledon Ground cover?/hygrophytic? Asteropollis asteroides CA Antophyta; Chloranthaceae Ground cover/hygrophytic (river banks, pioneer) Baculatisporites comaumensis SJ, CA, AP Monilophyta; Osmundaceae Ground cover/hygrophytic (riparian, mires, marshes) Baculatisporites spp. CA, AP Balmeiopsis limbata SJ, CA, AP Gymnospermophyta; Coniferales; Araucariaceae Canopy/xerophytic (coastal, upland?) Biretisporites potoniaei CA, AP Monilophyta; uncertain affinities Ground cover/hygrophytic (understory, wet lowland) Biretisporites spp. SJ, CA Botryococcus braunii SJ, CA, AP Chlorophyta; Botryococcaceae Brackish environments, fresh water bogs, temporary pools, ponds and lakes Brachyphyllum cf. obesum SJ Gymnospermophyta; Coniferales; Araucariaceae Canopy/xerophytic (coastal) ?Callaiosphaeridium assymetricum AP Dinoflagellata: Gonyaulacaceae Marine (Continued) PLOS ONE Late Albian desert biota from Iberia

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Table 1. (Continued) Taxon Section Known or probable parent affinity Possible preferred environment Callialasporites dampieri SJ, CA, AP Gymnospermophyta; Coniferales; Araucariaceae/Podocarpaceae Canopy/xerophytic (coastal, upland?) Callialasporites minor CA Callialasporites trilobatus CA, AP Callialasporites turbatus CA Callialasporites segmentatus AP Camarozonosporites ambigens CA Lycophyta; Lycopodiaceae Ground cover/hygrophytic (lowland, mires, wet environments) Camarozonosporites cf. ambigens CA Camarozonosporites cf. hammenii CA Camarozonosporites insignis AP Camarozonosporites spp. CA, AP Cardioangulina sp. CA Monilophyta; uncertain affinities Ground cover/hygrophytic? Cedripites cf. canadensis SJ Gymnospermophyta; Coniferales; Pinaceae Canopy/xerophytic (upland, well drained) Cedripites cf. cretaceous CA Cedripites sp. CA, AP Ceratosporites sp. SJ, CA Lycophyta; Selaginellaceae Ground cover/hygrophytic? Cerebropollenites macroverrucosus CA, AP Gymnospermophyta; uncertain affinity (taxodioid, Sciadopityaceae) Canopy/hygrophytic? (pioneer?) ?Chichaouadinium arabicum SJ Dinoflagellata: Peridiniaceae Marine Chlamydophorella sp. CA Dinoflagellata: uncertain affinity Marine Chomotriletes minor SJ, CA, AP Chlorophyta; Zygnemataceae Freshwaters Cibotiumspora jurienensis CA, AP Monilophyta; uncertain affinities Ground cover/hygrophytic (understory, wet lowland) Cibotiumspora sp. SJ Cicatricosisporites angicanalis CA Monilophyta; Schizaeales; Anemiaceae Ground cover, understory, epiphytes/ hygrophytic (lowland, marshes, wet

environments) Cicatricosisporites apicanalis SJ, CA, AP Cicatricosisporites aralicus SJ, AP Cicatricosisporites crassiterminatus CA Cicatricosisporites dorogensis CA Cicatricosisporites cf. grabowensis SJ Cicatricosisporites hughesii SJ, CA, AP Cicatricosisporites cf. ludbrookiae AP Cicatricosisporites cf. minutaestriatus AP Cicatricosisporites patapscoensis SJ, CA, AP Cicatricosisporites potomacensis SJ, CA, AP Cicatricosisporites proxiradiatus CA Cicatricosisporites pseudotripartitus CA Cicatricosisporites cf. pseudotripartitus SJ, CA, AP Cicatricosisporites cf. recticicatricosus CA Cicatricosisporites sinuosus CA, AP Cicatricosisporites sprumontii CA Cicatricosisporites venustus SJ, CA, AP Cicatricosisporites spp. SJ, CA, AP (Continued) PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 11 / 51

Table 1. (Continued) Taxon Section Known or probable parent affinity Possible preferred environment Cicatricosospoites auritus CA Monilophyta; Schizaeales; Anemiaceae Ground cover, understory, epiphytes/ hygrophytic (lowland, marshes, wet environments) Cicatricososporites phaseolus CA Cingutriletes clavus CA Bryophyta; Sphagnales; Sphagnaceae Ground cover/hygrophytic (marshes, peat bog?) Cingutriletes congruens SJ, CA Cingutriletes sp. CA, AP Circulodinium brevispinosum CA Dinoflagellata: Areoligeraceae Marine Classopollis major SJ, CA, AP Gymnospermophyta; Coniferales; Cheirolepidiaceae Canopy or shrubs/xerophytic (coastal, well drained lowland) Classopollis cf. obidosensis SJ, CA, AP Classopollis spp. SJ, CA, AP Clavatipollenites hughesii SJ, CA, AP Antophyta; Chloranthaceae Ground cover/hygrophytic (river banks, pioneer) Clavatipollenites tenellis SJ, CA, AP Clavatipollenites cf. tenellis CA Clavatipollenites spp. SJ, CA, AP Clavifera triplex CA Monilophyta; Gleicheniaceae Ground cover/xerophytic (lowland, pioneer) Clavifera sp. SJ, CA Cometodinium sp. CA Dinoflagellata: Gonyaulacaceae Marine Concavissimisporites cf. crassatus SJ, CA, AP Monilophyta; Dicksoniaceae/Lygodiaceae Ground cover/hygrophytic (understory, wet lowland) Concavissimisporites granulatus CA, AP Concavissimisporites cf. irregularis SJ, AP Concavissimisporites punctatus CA Concavissimisporites verrucosus CA Concavissimisporites spp. SJ, CA, AP Contignisporites cooksoniae SJ, AP Monilophyta; Pteridaceae Ground cover/hygrophytic (understory, wet

lowland) Contignisporites cf. cooksoniae CA Contignisporites sp. CA, AP Converrucosisporites geniculatus SJ, AP Monilophyta; uncertain affinities Ground cover/hygrophytic? Converrucosisporites spp. CA, AP Costatoperforosporites fistulosus CA, AP Monilophyta; Schizaeales; Anemiaceae Ground cover, understory, epiphytes/ hygrophytic (lowland, marshes, wet environments) Costatoperforosporites foveolatus SJ, CA, AP Costatoperforosporites triangulatus CA, AP Costatoperforosporites cf. triangulatus SJ Costatoperforosporites spp. SJ, CA, AP Couperisporites sp. AP Hepaticophyta Ground cover/xerophytic? Crassipollis chaloneri SJ, CA, AP Antophyta; uncertain affinity Ground cover?/hygrophytic? Crassipollis sp. CA Cretacaeiporites spp. SJ, CA, AP Antophyta; uncertain affinity (Trimeniaceae, Ranunculaceae, Eudicots) Ground cover?/hygrophytic? Cribroperidinium spp. AP Dinoflagellata: Gonyaulacaceae Marine (Continued) PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 12 / 51

Table 1. (Continued) Taxon Section Known or probable parent affinity Possible preferred environment Crybelosporites pannuceus SJ, CA, AP Monilophyta; Marsileaceae Ground cover/hygrophytic Cupuliferoidaepollenites parvulus CA Antophyta; non-magnoliid dicotyledon Canopy?/xerophytic? (lowland) Cupuliferoidaepollenites sp. SJ, CA, AP Cyathidites australis SJ, CA, AP Monilophyta; uncertain affinities Ground cover/hygrophytic (understory, wet lowland) Cyathidites minor SJ, CA, AP Cycadopites carpentieri CA Gymnospermophyta; uncertain affinity (Bennettitales, Cycadales, Czekanowskiales, Ginkgoales, Gnetales, Pentoxylales) Canopy/xerophytic (lowland, coastal) Cycadopites follicularis SJ, CA, AP Cycadopites spp. SJ, CA, AP Cyclonephelium chabacca CA, AP Dinoflagellata: Areoligeraceae Marine Cyclonephelium vannophorum CA Cyclonephelium spp. SJ, CA, AP Cymathiosphaera spp. CA, AP Chlorophyta (Prasinophyte) Marine Dammarites cf. albens AP Gymnospermophyta; Coniferales; Araucariaceae Canopy/xerophytic (coastal) Dammarites spp. SJ Dapsilidinium sp. SJ Dinoflagellata: uncertain affinity Marine Deltoidospora spp. SJ, CA, AP Monilophyta; uncertain affinities Ground cover/hygrophytic (understory, wet lowland) Densoisporites velatus CA Lycophyta; Pleuromeiaceae? Ground cover/hygrophytic (coastal, mires) Densoisporites sp. SJ, AP Dichastopollenites dunveganensis SJ, CA Antophyta; uncertain affinity Ground cover?, floating plant?/hygrophytic Dichastopollenites ghazalatensis CA Dichastopollenites cf. reticulatus AP Dichastopollenites spp. SJ, CA, AP

Dictyophyllidites equiexinus CA, AP Monilophyta; Dipteridaceae/Matoniaceae Ground cover/hygrophytic? Dictyophyllidites harrisii SJ, CA, AP Dictyophyllidites sp. CA Distachyapites sp. SJ Gymnospermophyta; Gnetales; Ephedraceae Shrubs?/xerophytic (lowland, coastal, upland) Distaltriangulisporites irregularis CA Monilophyta; Schizaeales; Schizaeaceae Ground cover/hygrophytic (understory, wet lowland) Distaltriangulisporites maximus CA Distaltriangulisporites cf. maximus CA Distaltriangulisporites perplexus SJ Distaltriangulisporites cf. perplexus CA Distaltriangulisporites sp. CA Distaverrusporites sp. AP Monilophyta; uncertain affinities Ground cover/hygrophytic? Doyleipollenites robbinsiae SJ, AP Antophyta; uncertain affinity Ground cover?/hygrophytic? Dryadopollis vestalis CA, AP Antophyta; non-magnoliid dicotyledon Ground cover?, shrubs?/xerophytic? (lowland) Echimonocolpites sp. AP Antophyta; uncertain affinity (Nymphaeaceae?, Monocots?) Floating plants/hygrohytic (mire, marshes) Echinatisporis varispinosus SJ Lycophyta; Selaginellaceae Ground cover, xerophytic? Echinatisporis sp. CA, AP Echinipollis cenomanensis CA Antophyta; uncertain affinity Ground cover?/xerophytic? (Continued) PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 13 / 51

Table 1. (Continued) Taxon Section Known or probable parent affinity Possible preferred environment Equisetosporites barghoornii CA Gymnospermophyta; Gnetales; Ephedraceae Shrub?/xerophytic (lowland, coastal, upland) Equisetosporites multicostatus CA, AP Equisetosporites aff. rousei SJ Equisetosporites spp. SJ, CA, AP Eretmophyllum sp. SJ Gymnospermophyta; Ginkgoales Canopy/xerophytic (lowland, marshes) Eucommiidites minor SJ, CA, AP Gymnospermophyta; Erdtmanithecales Canopy or shrub/xerophytic (lowland) Eucommiidites troedsonii SJ, CA, AP Eucommiidites spp. CA Exesipollenites tumulus SJ, CA, AP Gymnospermophyta; uncertain affinity (Bennettitales, taxodioid conifers) Canopy or shrub/xerophytic (coastal?, lowland?) Exesipollenites sp. SJ, CA, AP Exochosphaeridium sp. AP Dinoflagellata: uncertain affinity Marine Florentinia mantellii SJ, AP Dinoflagellata: Gonyaulacaceae Marine Florentinia sp. CA Foraminisporis asymmetricus SJ, CA, AP Hepaticophyta Ground cover, hygrophytic? Foraminisporis cf. undulatus CA Foraminisporites sp. CA Foveosporites cf. subtriangularis CA, AP Lycophyta Ground cover/hygrophytic (lowland, mires, wet environments) Foveotricolpites concinnus CA Antophyta; non-magnoliid dicotyledon Canopy?/xerophytic? (lowland) Foveotricolpites spp. CA, AP Foveotriletes cf. parviretus CA, AP Monilophyta; uncertain affinities Ground cover/hygrophytic? Fraxinoipollenites constrictus AP Antophyta; eudicots? Canopy?/xerophytic? (lowland) Fraxinoipollenites? sp. A sensu Barro´n et al. 2015 SJ, CA, AP Fraxinoipollenites? sp. B sensu

Barro´n et al., 2015 SJ Fraxinoipollenites? sp. AP Fraxinoipollenites spp. CA, AP Frenelopsis justae SJ Gymnospermophyta; Coniferales; Cheirolepidiaceae Canopy or shrubs/xerophytic (coastal, well drained lowland) Frenelopsis turolensis AP Frenelopsis cf. turolensis CA Ginginodinium cf. evittii SJ Dinoflagellata: Peridinaceae Marine Gleicheniidites cf. bolchovitinae SJ Monilophyta; Gleicheniaceae Ground cover/xerophytic (lowland, pioneer) Gleicheniidites carinatus CA Gleicheniidites laetus SJ Gleicheniidites senonicus SJ, CA, AP Gleicheniidites umbonatus CA Gnetaceaepollenites oreadis AP Gymnospermophyta; Gnetales; Ephedraceae Shrub/xerophytic (lowland, coastal, upland) Granulatisporites infirmus CA Monilophyta; Botryopteridales Ground cover/hygrophytic (wet lowland) Granulatisporites michinus CA Granulatisporites sp. CA, AP Gregussisporites orientalis SJ, CA, AP Monilophyta; uncertain affinities Ground cover/hygrophytic? (Continued) PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 14 / 51

Table 1. (Continued) Taxon Section Known or probable parent affinity Possible preferred environment Hammenia fredericksburgensis SJ, CA, AP Antophyta; Chloranthaceae Ground cover/hygrophytic (river banks, pioneer) Impardecispora cf. apiverrucata CA Monilophyta; Lygodiaceae Ground cover/hygrophytic (understory, wet lowland) Impletosphaeridium spp. SJ, CA, AP Dinoflagellata: uncertain affinity Marine Inaperturopollenites dubius SJ, CA, AP Gymnospermophyta; Coniferales; Cupressaceae (taxodioids) Canopy/hygrophytic (mire, marshes, river shores, wet lowland) Inaperturopollenites spp. SJ, CA, AP Gymnospermophyta; Coniferales; Araucariaceae?, Cupressaceae? Canopy/xerophytic? (lowland, coastal?) Ischyosporites crateris CA Monilophyta; Schizaeales; Schizaeaceae Ground cover/hygrophytic (understory, wet lowland) Ischyosporites cf. pseudoreticulatus AP Ischyosporites cf. punctatus CA Ischyosporites variegatus CA Ischyosporites sp. SJ, CA Jusinghipollis cf. ticoensis SJ, CA, AP Antophyta; uncertain affinity Ground cover?/hygrophytic?/canopy? ?Kallosphaeridium spp SJ Dinoflagellata: Gonyaulacaceae Marine Kiokansium unituberculatum CA, AP Dinoflagellata: uncertain affinity Marine Kraeuselisporites sp. (= Patellasporites? echinatus) SJ Lycophyta; Selaginellaceae? Ground cover, xerophytic? Kraeuselisporites spp. CA, AP Laevigatosporites ovatus CA, AP Monilophyta; uncertain affinities Ground cover/hygrophytic (lowland) Laevigatosporites spp. SJ, CA Leptolepidites cf. crassibalteus CA Unknown—Lycophyta?, Monilophyta? Ground cover/hygrophytic?

Leptolepidites proxigranulatus CA cf. Leptolepidites sp. SJ Leptolepidites spp. SJ, CA, AP Liliacidites cf. clavatus CA, AP Antophyta; probable monocot Ground cover/hygrophytic (wet lowland, marshes) Liliacidites cf. kiowaensis CA Liliacidites tectatus CA, AP Liliacidites spp. SJ, CA, AP “Liliacidites” minutus SJ, CA, AP Antophyta; Laurales Canopy?/xerophytic? (lowland) Lophotriletes sp. AP Monilophyta; Botryopteridales Ground cover/hygrophytic (wet lowland) ?Lusatisporites dettmannae SJ, CA, AP Monilophyta; uncertain affinities Ground cover/hygrophytic? Lycopodiumsporites sp. AP Lycophyta; Lycopodiaceae Ground cover/hygrophytic (lowland, mires, wet environments) cf. Margocolporites sp. SJ, CA Antophyta; Eudicots Canopy?/xerophytic? (lowland) Matonisporites crassiangulatus CA Monilophyta; Matoniaceae Ground cover/xerophytic? (lowland) Matonisporites sp. AP Micrhystridium sp. SJ, CA, AP Acritarcha: Acanthomorphitae Marine Microfoveolatosporis pseudoreticulatus CA Monilophyta; uncertain affinities Ground cover/hygrophytic Mirovia gothanii CA, AP Gymnospermophyta; Coniferales; Miroviaceae Canopy/xerophytic (lowland, marshes) Monocolpopollenites sp. SJ, CA, AP Antophyta; Arecales, Arecaceae Canopy/xerophytic? (lowland) (Continued) PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 15 / 51

Table 1. (Continued) Taxon Section Known or probable parent affinity Possible preferred environment Monosulcites minimus SJ, CA, AP Gymnospermophyta; uncertain affinity (Bennettitales, Cycadales, Ginkgoales) Canopy/xerophytic (lowland, coastal) Monosulcites sp. 1 CA, AP Monosulcites spp. SJ, CA, AP Montsechia-type SJ Antophyta; Ceratophyllum? Hygrophytic? Murospora sp. CA, AP Monilophyta; uncertain affinities Ground cover/hygrophytic? Nehvizdya (Eretmophyllum) penalveri CA, AP Gymnospermophyta; Ginkgoales Canopy/xerophytic (lowland, marshes) Neoraistrickia cf. densata CA Lycophyta; Selaginellaceae Ground cover, xerophytic? Neoraistrickia equalis CA Neoraistrickia robusta SJ, CA, AP Neoraistrickia truncata CA, AP Neoraistrickia spp. SJ, CA, AP Nevesisporites dailyi SJ, CA, AP Bryophyta Ground cover/hygrophytic (peat bog?) Nodosisporites segmentus AP Monilophyta; Schizaeales; Anemiaceae Ground cover, understory, epiphytes/ hygrophytic (lowland, marshes, wet environments) Nodosisporites sp. CA Odontochitina rhakodes SJ Dinoflagellata: Ceratiaceae Marine Oligosphaeridium complex SJ, CA, AP Dinoflagellata: Leptodinioideae Marine Oligosphaeridium spp. SJ, CA Osmundacidites wellmanii SJ, CA, AP Monilophyta; Osmundaceae Ground cover/hygrophytic (riparian, mire, marsh) Organic lining SJ Foraminifera Marine Ornamentifera peregrina SJ, CA Monilophyta; Gleicheniaceae Ground cover/xerophytic (lowland, pioneer) Ornamentifera cf. tuberculata CA Ornamentifera sp. CA, AP Ovoidinium sp. CA Dinoflagellata: Peridiniaceae Marine

Ovoidites sp. CA Chlorophyta; Zygnemataceae Freshwater Palaeohystrichophora sp. SJ Dinoflagellata: Peridiniaceae Marine Palaeoperidinium cretaceum SJ Parvisaccites amplus SJ Gymnospermophyta; uncertain affinity (Podocarpaceae, Voltziales) Canopy/xerophytic (upland, well drained) Parvisaccites radiatus CA, AP Parvisaccites spp. CA, AP aff. Parvisaccites spp. SJ Patellasporites tavaredensis SJ, CA, AP Monilophyta; uncertain affinities Ground cover/hygrophytic? Penetetrapites mollis CA Antophyta; uncertain affinity Canopy?/xerophytic? (lowland) Pennipollis escuchensis SJ, CA, AP Antophyta; Chloranthaceae-Ceratophyllum clade Ground cover/hygrophytic (wet lowland, marshes) Pennipollis cf. peroreticulatus SJ, CA, AP Pennipollis reticulatus SJ, CA, AP Pennipollis spp. SJ, CA, AP (Continued) PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 16 / 51

Reticulosporis gallicus CA Monilophyta; uncertain affinities Ground cover/hygrophytic (lowland) Retimonocolpites cf. crassatus CA Antophyta; uncertain affinity Ground cover?/hygrophytic? Retimonocolpites dividuus CA, AP Retimonocolpites cf. dividuus SJ Retimonocolpites excelsus CA Retimonocolpites cf. Rotundus SJ Retimonocolpites cf. textus SJ Retimonocolpites spp. SJ, CA, AP Retitrescolpites sp. A sensu Barro´n et al., 2015 SJ, AP Antophyta; non-magnoliid dicotyledon Canopy?/xerophytic? (lowland) Retitrescolpites spp. CA Retitriletes austroclavatidites CA, AP Lycophyta; Lycopodiaceae Ground cover/hygrophytic (lowland, mires, wet environments) Retitriletes clavatoides AP Retitriletes sp. SJ, CA, AP Rotverrusporites sp. CA Monilophyta; Osmundaceae Ground cover/hygrophytic (riparian, mire, marsh) Rousea georgensis SJ, CA, AP Antophyta; eudicots Canopy?/xerophytic? (lowland) Rousea cf. georgensis CA Rousea cf. delicata SJ Rousea miculipollis AP Rousea cf. prosimilis CA Rousea cf. scitula CA Rousea spp. CA, AP Ruffordiaspora sp. CA, AP Monilophyta; Schizaeales; Anemiaceae Ground cover, understory, epiphytes/ hygrophytic? (lowland, marshes) Rugubivesiculites sp. CA Gymnospermophyta; uncertain affinity Canopy/xerophytic (upland, well drained) (Continued) PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 17 / 51

Table 1. (Continued) Taxon Section Known or probable parent affinity Possible preferred environment Schizosporis reticulatus CA, AP Chlorophyta (Prasinophyte) Marine Sciadopityspollenites? sp. SJ, AP Gymnospermophyta; uncertain affinity (Sciadopityaceae?) Canopy/xerophytic (upland, well drained) Senectotetradites grossus CA Antophyta; eudicots; Nelumbonaceae? Floating plant?/hygrophytic (marshes) Senectotetradites spp. SJ, AP cf. Serialis sp. SJ Antophyta; uncertain affinity Hygrophytic? Sestrosporites pseudoalveolatus CA Lycophyta; Lycopodiaceae Ground cover/hygrophytic (lowland, mires, wet environments) Singhipollis microreticulatus SJ, CA Antophyta; uncertain affinity Ground cover?/hygrophytic? Spermatites sp. CA, AP Antophyta; uncertain affinity Hygrophytic? Spiniferites twistringiensis CA Dinoflagellata: Gonyaulacaceae Marine Spiniferites spp. SJ, CA Staplinisporites caminus SJ, AP Lycophyta; Lycopodiaceae Ground cover/hygrophytic (lowland, mires, wet environments) Staplinisporites cf. telatus AP Steevesipollenites sp. CA, AP Gymnospermophyta; Gnetales; Ephedraceae Shrub/xerophytic (lowland, coastal, upland) Stellatopollis cf. barghoornii SJ Antophyta; uncertain affinity (Magnoliids, Monocots) Groun cover/hygrophytic (pioneer, mire, marshes) Stellatopollis sp. AP Stereisporites psilatus SJ, CA Bryophyta; Sphagnales; Sphagnaceae Ground cover/hygrophytic (marshes, peat bog?) Stereisporites spp. SJ, CA, AP Striatella cf. balmei CA Monilophyta; uncertain affinities Ground cover/hygrophytic Striatopollis sp. SJ, CA, AP

Antophyta; Eudicots Canopy?/xerophytic? (lowland) Subtilisphaera sp. CA Dinoflagellata: Peridiniaceae Marine Tasmanaceae? Gen. et sp. indet. AP Chlorophyta (Prasinophyte) Marine Tasmanites sp. CA Chlorophyta (Prasinophyte) Marine Taurocusporites segmentatus SJ, CA, AP Bryophyta; Sphagnales; Sphagnaceae Ground cover/hygrophytic (marshes, peat bog?) Taurocusporites sp. CA, AP Taxodiaceaepollenites hiatus SJ, CA, AP Gymnospermophyta; Coniferales; Cupressaceae (taxodioids) Canopy/hygrophytic (mire, marshes, river shores, wet lowland) Tehamadinium sp. CA Dinoflagellata: Leptodinioideae Marine Tenua hystrix SJ Dinoflagellata: Gonyaulacaceae Marine Tetracolpites sp. CA Antophyta; Eudicots Canopy?/xerophytic? (lowland) Tetraporina sp. SJ Chlorophyta; Zygnemataceae Freshwaters Tigrisporites scurrandus SJ Monilophyta; Filicales Ground cover/hygrophytic (lowland) Tigrisporites sp. CA Transitoripollis anulisulcatus SJ, CA, AP Antophyta; Chloranthaceae-Ceratophyllum clade Groun cover?/hygrophytic? Transitoripollis similis CA, AP Transitoripollis cf. similis SJ Transitoripollis spp. SJ, CA, AP Trichodinium castanea CA, AP Dinoflagellata: Gonyaulacaceae Marine (Continued) PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 18 / 51

Table 1. (Continued) Taxon Section Known or probable parent affinity Possible preferred environment Tricolpites cf. amplifissus AP Antophyta; eudicots Canopy?/xerophytic? (lowland) Tricolpites cf. brnicensis AP Tricolpites cf. maximus CA, AP Tricolpites micromunus SJ, CA, AP Tricolpites minutus CA, AP Tricolpites cf. minutus SJ Tricolpites nemejci SJ, CA, AP Tricolpites cf. parvus SJ, CA, AP Tricolpites sagax CA, AP Tricolpites cf. sagax CA Tricolpites vulgaris AP Tricolpites cf. vulgaris SJ, CA, AP Tricolpites sp. cf. Retitricolpites varireticulatus SJ, CA Tricolpites spp. SJ, CA, AP Tricolporoidites cf. pacltovae CA Antophyta; non-magnoliid dicotyledon Canopy?/xerophytic? (lowland) Tricolporoidites spp. CA, AP Trilobosporitesspp. SJ, CA, AP Monilophyta; Lygodiaceae Ground cover/hygrophytic (understory, wet lowland) Triporoletes cenomanianus CA Hepaticophyta; cf. Ricciaceae Ground cover/hydrophytic? Triporoletes radiatus CA Triporoletes reticulatus CA Triporoletes simplex AP Triporoletes sp. SJ, CA Tucanopollis crisopolensis SJ, CA Tucanopollis spp. CA Antophyta; Chloranthaceae-Ceratophyllumclade Groun cover?/hygrophytic? Uesuguipollenites callosus SJ, CA, AP Gymnospermophyta; Coniferales; Araucariaceae Canopy/xerophytic (coastal, upland?) Undulatisporites cf. pflugii CA Monilophyta; uncertain affinities Ground cover/hygrophytic Undulatisporites cf. rugulatus AP Undulatisporites undulapolus CA Undulatisporites spp. SJ, CA, AP Uvaesporites sp. CA Lycophyta; Selaginellaceae Ground cover/riparian?, xerophytic?

Varirugosisporites tolmanensis CA Monilophyta; uncertain affinities Ground cover/hygrophytic Varirugosisporites sp. CA Verrucatosporites sp. CA, AP Monilophyta; uncertain affinities Ground cover/hygrophytic Verrucosisporites cf. rotundus CA, AP Monilophyta; uncertain affinities Ground cover/hygrophytic aff. Verrucosisporites kopukuensis CA, AP Verrucosisporites spp. CA, AP Virgo amiantopollis CA, AP Antophyta; Eudicots (Gunnerales) Groun cover?/hygrophytic? Vitreisporites pallidus SJ, CA, AP Pteridospermatophyta; Caytoniales Canopy/hygrophytic (lowland, deltaic, mire, floodplains) Widdringtonites sp. SJ Gymnospermophyta; Coniferales; Cupressaceae Canopy/xerophytic (lowland) https://doi.org/10.1371/journal.pone.0282178.t001 PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 19 / 51

Table 2. List of arthropod taxa as bioinclusions identified from San Just, Arroyo de la Pascueta and La Hoya Cretaceous amber-bearing outcrops with information about autoecology of each species and possible preferred environment. Arthropod taxon Family Order Amber outcrop Autoecology, preferred environment Actenobius magneoculus Ptinidae Coleoptera San Just Wood-borer on conifers? Alavaromma orchamum Alavarommatidae Hymenoptera San Just Parasitoid? Ametroproctus valeriae Ametroproctidae Acariformes San Just Detritivorous, tree bark habitat Aragomantispa lacerata Mantispidae Neuroptera San Just Predator Aragonimantis aenigma Incertae sedis Mantodea San Just Predator Aragonitermes teruelensis Incertae sedis Blattodea: Isoptera San Just Recycler of lignocellulose Archaeatropos alavensis Empheriidae Psocodea San Just and Arroyo de la Pascueta Detritivorous, decayed leaf-litter habitat Archaeognatha gen. et sp. indet. Archaeognatha San Just Detritivorous, decayed leaf-litter habitat Archiculicoides skalskii Ceratopogonidae Diptera San Just Blood-feeder Arra legalovi Nemonychidae Coleoptera San Just Pollen-feeder on Araucariaceae? Blattodea gen. et sp. indet. Blattodea San Just, Arroyo de la Pascueta and La Hoya Omnivorous, decayed leaf-litter habitat Burmaphron jentilak Stigmaphronidae Hymenoptera San Just Parasitoid? Burmazelmira grimaldii Archizelmiridae Diptera San Just Unknown Collembola gen. et sp. indet. Collembola San Just Detritivorous, decayed leaf-litter habitat

Cretaceobodes martinezae Otocepheidae Acariformes San Just Detritivorous, decayed leaf-litter habitat Cretaceomma turolensis Gallorommatidae Hymenoptera San Just Parasitoid? Cretevania alcalai Evaniidae Hymenoptera San Just Parasitoid Cretevania montoyai Evaniidae Hymenoptera San Just Parasitoid Cretevania rubusensis Evaniidae Hymenoptera Arroyo de la Pascueta Parasitoid Diameneura marveni Spathiopterygidae Hymenoptera San Just Parasitoid? Diptera gen. et sp. indet. Diptera San Just, Arroyo de la Pascueta and La Hoya Diverse autoecology Hispanothrips utrillensis Stenurothripidae Thysanoptera San Just Herbivorous Helius turolensis Limoniidae Diptera San Just Pollen-feeder? Hemiptera gen. et sp. indet. Hemiptera San Just and Arroyo de la Pascueta Phytophagous Hymenoptera gen. et sp. indet. Hymenoptera San Just, Arroyo de la Pascueta and La Hoya Diverse autoecology Hypovertex hispanicus Scutoverticidae Acariformes San Just Detritivorous, decayed leaf-litter habitat Iberofoveopsis miguelesi Perforissidae Hemiptera San Just Phytophagous, wet environments Leptoconops zherikhini Ceratopogonidae Diptera San Just Blood-feeder Leptus sp. Erythraeidae Acariformes San Just Parasitoid Litoleptis fossilis Rhagionidae Diptera San Just Blood-feeder? Microphorites utrillensis Dolichopodidae Diptera San Just Predator, sandy beach environment Mymaropsis turolensis Spathiopterygidae Hymenoptera San Just Parasitoid? Orchestina sp. Oonopidae Araneae San Just Predator