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

Comparison with other localities The palynological assemblages from the Maestrazgo Basin documented here are similar to coe- val successions from the Las Parras and Oliete sub-basins [31, 33–35]. Afropollis (Fig 7J), ephe- droid polyplicate pollen (Fig 7O) and several angiosperm taxa such as Dichastopollenites (Fig 9G–9I and 9L) and Cretacaeiporites relate the Spanish assemblages to the palynofloras from North Gondwana (see i.e. [210–214]) and support Iberia as a transitional zone between the flo- ral provinces of NE Gondwana and Laurasia [104]. The Maestrazgo palynological assemblages can also be related to the ones described from the Basque-Cantabrian Basin [31, 87, 104, 132] as both basins share more than 90% of the identified miospore families and a significant num- ber of their respective species. The faunal record, particularly the arthropod assemblages char- acterising the two Iberian areas, reveals a series of common species [162]. Arthropods recorded in the Maestrazgo with a more widespread distribution include Cretevania (Hyme- noptera: Evaniidae) documented in various outcrops across Europe and Asia [215], Serphites (Hymenoptera: Serphitidae) recorded in Europe, Asia, and North America [216] and Micro- phorites (Diptera: Dolichopodidae) found in ambers from France, Lebanon and Myanmar [168]. The coeval miospore assemblages from the Lusitanian Basin exhibit higher angiosperm abundance than the ones described herein [6, 105, 113, 217]. While this difference could likely reflect vegetation with more angiosperms, the higher representation of dinocysts of the Portu-

guese successions suggests the influence of the depositional environments as sedimentary suc- cessions on a passive margin can collect miospores from a larger catchment area. The megafloral record relates the flora from the Maestrazgo to Laurasian assemblages. Tax- onomically similar, low-diversity plant assemblages have been documented from mid-Creta- ceous strata of southwestern and northern France [218–220] and the Czech Republic [83, 172]. These shared characteristics are interesting as they suggest that a few species of angiosperms were thriving in salt-influenced environments and that geographic and climatic barriers did not prevent a widespread distribution across Laurasia. Pathways allowing the dispersal of taxa with such specialised ecological requirements could include coastal fringes surrounding the Tethys. The good representation of Tucanopollis in both NE Gondwana and southern Laurasia suggests that a significant number of halophytic angiosperms used this ‘coastal pathway’ to col- onise new environments. However, more direct, NS trending migration routes following northern and southern Atlantic rift valleys could also be considered to link central Gondwana floras of Gabon and Brazil to Laurasian floras. Despite the numerous similarities existing between floras from Gondwana, mid and north- ern Laurasia, it is probable that the latitudinal migration of flora remained hindered by differ- ences in climate. Floras from locations at equivalent palaeolatitudes, should always present

more compositional and structural similarities than floras from distinct climatic belts. Late Albian–early Cenomanian floras from NW Europe and N America present higher percentages in Gleicheniaceae and Cyathidites but lower diversity in angiosperms than coetaneous floras from southern Europe. Latitudinal effects influenced the composition of the mid-Cretaceous palynological assem- blages from Western Europe and the Atlantic Coastal Plain of USA, since they belonged to the Northern Mid-latitude Warm humid belt (NMW belt) [17, 221]. Although the diversity of taxa was high in the palynological assemblages attributed to the NMW belt, the abundance of mios- pores is different from that of the Maestrazgo Basin (i.e., [90, 112, 222–226]), which was under the influence of the Northern Hot Arid (NHA) belt [17, 188] during the late Albian. More con- cretely, under the influence of the NMW belt, fern spores were generally abundant, showing conspicuous amounts of Gleicheniaceae and Cyathidites, the amounts of Classopollis indicate PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 39 / 51

that it was a sub-dominant pollen type, bisaccate and inaperturate conifer pollen reaches con- spicuous values, and angiosperm pollen exhibits high diversity and significant abundances (i.e., abundances of more than 30% of angiosperm pollen, and a diversity of around 47 taxa in Zone II of the Potomac Group [112]). Additionally, the late Albian–early Cenomanian palynofloras from the Northern High-lati- tude Temperate Humid (NHT) belt are characterised by a high diversity of dinocysts, trilete spores and angiosperms (see i.e., [111, 227–229]). For example, the coeval existence of quite different plant communities from those described in eastern Iberia can be regarded if compar- ing the late Albian floras from Peace River Area (Alberta, Canada) [229]. These floras are char- acterised by scarce Classopollis, high amounts of bisaccate pollen grains of the genera Alisporites and Vitreisporites, trilete spores such as Cingutriletes, Gleicheniidites, and Stereispor- ites, as well as conspicuous values of tricolpate angiosperm species (Cupuliferoidaepollenites parvulus, Tricolpites sagax, T. vulgaris). Conclusions The palaeobotanical study of three sections from the Utrillas Group in the Maestrazgo Basin (Iberian Ranges, eastern Spain) reveals the existence of ancient conifer woodlands and fern/ angiosperm communities that thrived in the mid-Cretaceous Iberian Desert System. Four veg- etation types have been inferred: the first one is characterised by arid woodlands of Cheirolepi- diaceae-Cupressaceae, the second is related to coastal settings with Araucariaceae, the third is

related to salt marshes, and the fourth represents local fern and angiosperm communities linked to wet interdunes (oases) and wetlands. The existence of ponded areas linked to wet interdunes is interpreted based on the high percentages of fern spores, the common occur- rence of angiosperm pollen and the lack of marine palynomorphs. The presence of spikes of Tucanopollis spp. and “Liliacidites” minutus in the CA section might be related to the relevant role of the angiosperms in the colonisation of xeric and disturbed habitats and the ecological successions in arid settings. The biostratigraphic markers recorded in the studied sections confirm a late Albian age, except for the uppermost levels of the Cortes de Arenoso section (La Hoya amber outcrop), in which an early Cenomanian age might be inferred based on the occurrence of Echinipollis cen- omanensis, Polypodiisporonites cenomanianus and Senectotetradites grossus. Although the late Albian palynoflora from the Maestrazgo Basin was influenced by the palaeoclimatic conditions that prevailed in the Northern Hot Arid belt, the studied assemblages can generally be related to others from Europe and North America. However, the presence of pollen grains of the gen- era Afropollis, Equisetosporites, Gnetaceapollenites, Steevesipollenites, Dichastopollenites and Cretacaeiporites indicates that Iberia was a transitional area between the Northern Gondwanan and the Southern Laurasian provinces. In this sense, the studied macrofloristic assemblages from the Maestrazgo Basin show similarities to others from the European Laurasia.

Supporting information S1 Appendix. Absolute abundances of palynomorphs in the studied sections from the Maestrazgo Basin. (XLSX) S2 Appendix. List of the identified taxa ordered according their botanical and zoological identification, and the alphabetic citation of the taxonomic authorities. (DOC) PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 40 / 51

Acknowledgments The authors would like to thank Dr Rafael Lo´pez del Valle for the preparation of the amber pieces with arthropod specimens, Dr Juan Pedro Rodrı´guez-Lo´pez for his assistance in the fieldwork, Pepa Torres Matilla for her aid with the design of the figures, Dr Xavier Delclòs as supervisor of the PhD Thesis of SA´ P, Dr Luis Somoza (CN-IGME CSIC) for the financial sup- port (AYUDAS EXTRAORDINARIAS MENCIONES EXCELENCIA SEVERO OCHOA del IGME-CSIC) and the Direccio´n General de Patrimonio Cultural del Gobierno de Arago´n (Spain) for granting the permissions of excavation. We would also like thank the editor (Dr Enrique Peñalver) and the two anonymous referees who provided valuable suggestions for improving the quality of the manuscript. Author Contributions Conceptualization: Eduardo Barro´n, Daniel Peyrot. Data curation: Eduardo Barro´n. Formal analysis: Eduardo Barro´n, Daniel Peyrot. Funding acquisition: Eduardo Barro´n. Investigation: Eduardo Barro´n, Daniel Peyrot, Carlos A. Bueno-Cebollada, Jiřı´ Kvaček, Sergio A´ lvarez-Parra, Yul Altolaguirre, Nieves Mele´ndez. Methodology: Eduardo Barro´n, Daniel Peyrot, Jiřı´ Kvaček, Sergio A´ lvarez-Parra, Nieves Mele´ndez. Resources: Eduardo Barro´n. Supervision: Eduardo Barro´n. Visualization: Daniel Peyrot, Carlos A. Bueno-Cebollada, Jiřı´ Kvaček, Sergio A´ lvarez-Parra, Yul Altolaguirre. Writing – original draft: Eduardo Barro´n, Daniel Peyrot, Carlos A. Bueno-Cebollada. Writing – review & editing: Eduardo Barro´n, Daniel Peyrot, Carlos A. Bueno-Cebollada. References

1. Perrichot V. Environnements paraliques à ambre et à ve´ge´taux du Cre´tace´ Nord-Aquitain (Charentes, Sud-Ouest de la France). PhD Thesis. Univ Rennes. 2003. 2. Peyrot D, Rodrı´guez-Lo´pez JP, Barro´ n E, Mele´ndez N. Palynology and biostratigraphy of the Escucha Formation in the Early Cretaceous Oliete Sub-basin, Teruel, Spain. Rev Esp Micropaleontol. 2007; 39 (1–2): 135–154. 3. Die´ guez C, Peyrot D, Barro´ n E. Floristic and vegetational changes in the Iberian Peninsula during Jurassic and Cretaceous. Rev Palaeobot Palynol. 2010; 162: 325–340. 4. Coiffard C, Gomez B, Daviero-Gomez V, Dilcher D. Rise to dominance of angiosperm pioneers in European Cretaceous environments. PNAS. 2012; 109(51): 20955–20959. https://doi.org/10.1073/ pnas.1218633110 PMID: 23213256 5. Peralta-Medina E, Falcon-Lang HJ. Cretaceous forest composition and productivity inferred from a global fossil wood database. Geology. 2012; 40 (3): 219–222. 6. Heimhofer U, Hochuli PA, Burlas S, Oberli F, Adatte T, Dinis JL, et al. Climate and vegetation history of western Portugal inferred from Albian near-shore deposits (Gale´ Formation, Lusitanian Basin). Geol Mag. 2012; 149: 1046–1064. 7. Heimhofer U, Wucherpfenning N, Adatte T, Schouten S, Schneebeli-Hermann E, Gardin S, et al. Veg- etation response to exceptional global warmth during Oceanic Anoxic Event 2. Nat Commun. 2018; 9: 3832. https://doi.org/10.1038/s41467-018-06319-6 PMID: 30237441 PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 41 / 51

8. Vakhrameev VA. Jurassic and Cretaceous floras and climates of the Earth. Cambridge: Cambridge Univ Press; 1991. 9. Herngreen GFW, Kedves M, Rovnina LV, Smirnova SB. Chapter 29C. Cretaceous palynofloral prov- inces: a review. In: Jansonius J, McGregor DC, editors. Palynology: Principles and Applications, vol. 3. Salt Lake City: Am Assoc Stratigr Palynol Found; 1996. pp. 1157–1188. 10. Lidgard S, Crane PR. Angiosperm diversification and Cretaceous floristic trends: a comparison of paly- nofloras and leaf macrofloras. Paleobiology. 1990; 16: 77–93. 11. Boyce CK, Brodribb TJ, Field TS, Zwieniecki MA. Angiosperm leaf vein evolution was physiologically and environmentally transformative. Proc R Soc Lond B. 2009; 276: 1771–1776. https://doi.org/10. 1098/rspb.2008.1919 PMID: 19324775 12. Friis EM, Crane PR, Pedersen KR. Early flowers and angiosperm evolution. Cambridge: Cambridge Univ Press; 2011. 13. Herendeen PS, Friis EM, Perdersen KR, Crane PR. Palaeobotanical redux: revisiting the age of the angiosperms. Nat Plants. 2017; 17015. https://doi.org/10.1038/nplants.2017.15 PMID: 28260783 14. Rodrı´guez-Lo´pez JP, Peyrot D, Barro´ n E. Complex sedimentology and palaeohabitats of Holocene coastal deserts, their topographic controls, and analogues for the mid-Cretaceous of northern Iberia. Earth Sci Rev. 2020; 201: 103075. 15. Coiro M, Doyle JA, Hilton J. How deep is the conflict between molecular and fossil evidence on age of angiosperms? New Phytol. 2019; 223(1): 83–99. 16. Boukhamsin H, Peyrot D, Vecoli M. Angiosperm pollen assemblages from the Lower Cretaceous (Bar-

24. Ch Wu, Ch Kiu, yi H, Xia G, Zhang H, Wang L, et al. Mid-Cretaceous desert system in the Simao Basin, southwestern China, and its implications for sea-level change during a greenhouse climate. Palaeogeogr Palaeoclimatol Palaeoecol. 2017; 468: 529–544. 25. Bueno-Cebollada CA, Barro´n E, Peyrot D, Mele´ ndez N. Palynostratigraphy and palaeoenvironmental evolution of the Aptian to lower Cenomanian succession in the Serranı´a de Cuenca (Eastern Spain). Cretaceous Res. 2021; 128: 104956. 26. Smith RMH, Mason TR. Sedimentary environments and trace fossils of Tertiary oasis deposits in the Central Namib Desert, Namibia. Palaios. 1998; 13: 547–559. 27. Parris JT, Falcon-Lang HJ. Coniferous trees associated with interdune deposits in the Jurassic Navajo Sandstone Formation, Utah, USA. Palaeontology. 2007; 50: 829–843. 28. Gomez B, Barale G, Martı´n-Closas C, The´venard F, Philippe M. De´couverte d’une flore à Ginkgoales, Bennettitales et Conife´ rales dans le Cre´ tace´ infe´ rieur de la Formation Escucha (Chaıˆne Ibe´rique Orientale, Teruel, Espagne). Neues Jahr Geol Pala¨ ontol Monatsh. 1999; 1999(11): 661–675. 29. Gomez B, Martı´n-Closas C, Barale G, The´venard F. A new species of Nehvizdya (Ginkgoales) from the Lower Cretaceous of the Iberian Ranges (Spain). Rev Palaeobot Palynol. 2000; 111: 49–70. https://doi.org/10.1016/s0034-6667(00)00017-8 PMID: 10936606 30. Gomez B. A new species of Mirovia (Coniferales, Miroviaceae) from the Lower Cretaceous of the Ibe- rian Ranges (Spain). Cretaceous Res. 2002; 23: 761–773. PLOS ONE Late Albian desert biota from Iberia

PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 42 / 51

31. Villanueva-Amadoz U. Nuevas aportaciones palinostratigra´ ficas para el intervalo Albiense-Cenoma- niense en el Sector NE de la penı´nsula Ibe´rica. Implicaciones paleogeogra´ficas y paleoclima´ ticas. PhD Thesis. Univ Zaragoza. 2009. 32. Villanueva-Amadoz U, Pons D, Dı´ez JB, Ferrer J, Sender LM. Angiosperm pollen grains of Sant Just site (Escucha Formation) from the Albian of the Iberian Range (north-eastern Spain). Rev Palaeobot Palynol. 2010; 162(3): 362–381. 33. Villanueva-Amadoz U, Sender LM, Dı´ez JB, Ferrer J, Pons D. Palynological studies of the boundary marls unit (Albian–Cenomanian) from northeastern Spain. Paleophytogeographical implications. Geo- diversitas. 2011; 33(1): 137–176. 34. Sender LM, Gomez B, Dı´ez JB, Coiffard C, Martı´n-Closas C, Villanueva-Amadoz U, et al. Ploufolia cerciforme gen. et comb. nov.: Aquatic angiosperm leaves from the Upper Albian of north-eastern Spain. Rev Palaeobot Palynol. 2010; 161: 77–86. 35. Sender LM, Villanueva-Amadoz U, Dı´ez JB, Sa´ nchez-Pellicer R, Bercovici A, Pons D, et al. A new uppermost Albian flora from Teruel province, northeastern Spain. Geodiversitas. 2012; 34(2): 373– 397. 36. Sender LM, Doyle JA, Upchurch GR, Villanueva-Amadoz U, Dı´ez JB. Leaf and inflorescence evidence for near-basal Araceae and an unexpected diversity of other monocots from the late Early Cretaceous of Spain. J Syst Palaeontol. 2019; 17(15): 1093–1126. 37. Barral A, Gomez G, Daviero-Gomez V, Le´ cuyer C, Mendes MM, Ewin TAM. New insights into the mor-

phology and taxonomy of the Cretaceous conifer Frenelopsis based on a new species from the Albian of San Just, Teruel, Spain. Cretaceous Res. 2019; 96: 21–36. 38. Vilanova y Piera J. Manual de Geologı´a aplicada a la Agricultura y a las Artes Industriales, vol. 1. Madrid: Imprenta Nacional; 1860. 39. Delclòs X, Arillo A, Peñalver E, Barro´n E, Soriano C, Lo´ pez del Valle R, et al. Fossiliferous amber deposits from the Cretaceous (Albian) of Spain. CR Palevol. 2007; 6: 135–149. 40. Peñalver E, Delclòs X. Spanish amber. In: Penney D., editor. Biodiversity of fossils in amber from the major world deposits. Manchester: Siri Scientific Press; 2010. pp. 236–270. 41. A´ lvarez-Parra S, Pe´rez-de la Fuente R, Peñalver E, Barro´n E, Alcala´ L, Pe´ rez-Cano J, et al. Dinosaur bonebed amber from an original swamp forest soil. eLife. 2021; 10:e72477. https://doi.org/10.7554/ eLife.72477 PMID: 34844669 42. Peñalver E, Martı´nez-Delclòs X. Importancia patrimonial de Arroyo de la Pascueta, un yacimiento de a´mbar creta´ cico con insectos fo´ siles en Rubielos de Mora. In: Mele´ ndez Hevia G, Peñalver Molla´ E, editors. El patrimonio paleontolo´gico de Teruel. Teruel: Instituto de Estudios Turolenses; 2002. pp. 201–208. 43. Peñalver E, Delclòs X, Soriano C. A new rich outcrop with palaeobiological inclusions in the Lower Cretaceous of Spain. Cretaceous Res. 2007; 28: 791–802. 44. Peñalver E, Ortega-Blanco J, Nel A, Delclòs X. Mesozoic Evaniidae (Insecta: Hymenoptera) in Span- ish amber: reanalysis of the phylogeny of the Evanioidea. Acta Geol Sin. 2010; 84 (4): 809–827.

45. Aurell M, Ba´denas B, Gasca JM, Canudo JI, Liesa CL, Soria AR, et al. Stratigraphy and evolution of the Galve sub-basin (Spain) in the middle Tithonian–early Barremian: implications for the setting and age of some dinosaur fossil sites. Cretaceous Res. 2016; 65: 138–162. 46. Cane´ rot J, Crespo Zamorano A, Navarro Va´zquez D. Hoja geolo´ gica n˚ 518 (Montalba´n). Mapa Geolo´- gico de España 1:50.000. 2ª serie. Madrid: IGME; 1977. 47. Almera J, Anado´ n P, Godoy A. Mapa Geolo´ gico de España 1:50.000, hoja n˚ 591 (Mora de Rubielos). Madrid: IGME; 1972. 48. Sopeña A, Gutie´rrez-Marco JC, Sa´ nchez-Moya Y, Go´mez JJ, Mas R, Garcı´a A, et al. Cordillera Ibe´ rica y Costero Catalana. In: Vera JA, editor. Geologı´a de España. Madrid: SGE-IGME; 2004. pp. 465–527. 49. Martı´n-Chivelet J, Lo´pez-Go´mez J, Aguado R, Arias C, Arribas J, Arribas ME, et al. The Late Juras- sic–Early Cretaceous rifting. In: Quesada C, Oliveira JT, editors. The Geology of Iberia: A Geodynamic Approach. Volume 3: The Alpine cycle. Cham: Springer; 2019. pp. 169–249. 50. Cane´ rot J, Cugny P, Pardo G, Salas R, Villena J. Ibe´ rica central-maestrazgo. In: Garcı´a A, editor. El Creta´ cico de España. Madrid: Univ Complutense; 1982. pp. 273–344. 51. Soria AR. La sedimentacio´n en las cuencas marginales del Surco Ibe´ rico durante el Creta´ cico Inferior y su control estructural. PhD Thesis. Univ Zaragoza. 1997. 52. Salas R, Guimerà J, Mas R, Martı´n-Closas C, Mele´ ndez A, Alonso A. Evolution of the Mesozoic Cen-

tral Iberian Rift System and its Cainozoic inversion (Iberian chain). Peri-Tethys Mem. 2001; 6: 145– 185. PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 43 / 51

53. Peropadre C. El Aptiense del margen occidental de la Cuenca del Maestrazgo: controles tecto´ nico, eusta´ tico y clima´tico en la sedimentacio´n. PhD Thesis. Madrid: Univ Complutense. 2011. 54. Peropadre C, Liesa CL, Mele´ndez N. High-frequency, moderate to high-amplitude sea-level oscilla- tions during the late early Aptian: Insights into the mid-Aptian event (Galve Sub-basin, Spain). Sedi- ment Geol. 2013; 294: 233–250. 55. Aurell M, Fregenal-Martı´nez M, Ba´denas B, Muñoz-Garcı´a MB, E´ lez J, Mele´ ndez N, et al. Middle Jurassic–Early Cretaceous tectono-sedimentary evolution of the southwestern Iberian Basin (central Spain): Major palaeogeographical changes in the geotectonic framework of the Western Tethys. Earth Sci Rev. 2019; 199: 1–33. 56. Cervera A, Pardo G, Villena J. Algunas precisiones litoestratigra´ ficas sobre la formacio´ n «Lignitos de Escucha». Tecniterrae. 1976; 14: 25–33. 57. Pardo G. Estratigrafı´a y sedimentologı´a de las formaciones detrı´ticas del Creta´ cico Inferior terminal del Bajo Arago´ n Turolense. PhD Thesis. Univ Zaragoza. 1979. 58. Querol X, Salas R, Pardo G, Ardevol L. Albian coal-bearing deposits of the Iberian Range in northeast- ern Spain. In: Mc. Cabe JP, Panish JT, editors. Coals. Geol Soc Am Spec Pap. 1992; 267: 193–208. 59. Rodrı´guez-Lo´pez JP, Mele´ ndez N, Soria AR, De Boer P. Reinterpretacio´ n estratigra´ fica y sedimento- lo´gica de las Formaciones Escucha y Utrillas de la Cordillera Ibe´ rica. Rev Soc Geol Esp. 2009; 22(3– 4): 163–219.

60. Rodrı´guez-Lo´pez JP. Sedimentologı´a y evolucio´n del sistema dese´ rtico arenoso (erg) desarrollado en el margen occidental del Tethys durante el Creta´cico Medio, Cordillera Ibe´rica. Provincias de Teruel y Zaragoza. PhD Thesis. Madrid: Univ Complutense. 2008. 61. Aguilar MJ, Ramı´rez del Pozo J, Riba O. Algunas precisiones sobre la sedimentacio´ n y paleocecolo- gı´a del Creta´ cico Inferior en la zona de Utrillas-Villarroya de los Pinares (Teruel). Estud Geol. 1971; 27(6): 497–512. 62. Pardo G, Ardevol L, Villena J. Mapa Geolo´ gico de España. 1:200.000, hoja n˚40 (Daroca). Madrid: ITGE; 1991. 63. Liesa CL, Casas AM, Soria AR, Simo´ n JL, Mele´ ndez A. Estructura extensional creta´ cica e inversio´ n terciaria en la regio´n de Aliaga-Montalba´ n. Geo-Guı´as. 2004; 1: 151–180. 64. Martı´n-Chivelet J, Floquet M, Garcı´a-Senz J, Callapez PM, Lo´ pez-Mir B, Muñoz JA, et al. Late Creta- ceous Post-Rift to Convergence in Iberia. In: Quesada C, Oliveira JT, editors. The Geology of Iberia: A Geodynamic Approach. Volume 3: The Alpine cycle. Cham: Springer; 2019. pp. 285–376. 65. Altolaguirre Y. Estudio palinolo´gico preliminar del Creta´cico de Cortes de Arenoso, Castello´ n. MSc Thesis. Madrid: Univ Complutense; 2015. 66. Zervas D, Nichols GJ, Hall R, Smyth HR, Lu¨ thje C, Murtagh F. SedLog: A shareware program for drawing graphic logs and log data manipulation. Comput Geosci. 2009; 35 (10): 2151–2159. 67. Batten DJ. Small palynomorphs. In: Jones TP, Rowe NP, editors. Fossil plant and spores modern techniques. London: Geol Soc; 1999. pp. 15–19.

68. Paleopalynology Traverse A., Second ed. Dordrecht: Springer; 2007. 69. Grimm EC. Tilia, Version 2. Springfield: Illinois Stat Mus; 1992. 70. Grimm EC. TGView, Version 2.0.2. Springfield: Illinois Stat Mus; 2004. 71. Jansonius J, McGregor DC. Palynology: principles and applications. Volume 1. Principles. Salt Lake City: Am Assoc Stratigr Paynol Found; 1996. 72. Peyrot D. Late Cretaceous (late Cenomanian-early Turonian) dinoflagellate cysts from the Castilian Platform, northern Spain. Palynology. 2011; 35(2): 267–300. 73. Van Konijnenburg-Van Cittert JHA. Ecology of some Late Triassic to Early Cretaceous ferns in Eur- asia. Rev Palaeobot Palynol. 2002; 119: 113–124. 74. Abbink OA, Van Konijnenburg-Van Cittert JHA, Visscher H. A sporomorph ecogroup model for the Northwest European Jurassic–Lower Cretaceous: concepts and framework. Geol Mijnbouw. 2004; 83: 17–31. 75. Mohr BAR, Bernardes-de-Oliveira MEC, Loveridge R, Pons D, Sucerquia PA, Castro-Fernandes MC. Ruffordia goeppertii (Schizaeales, Anemiaceae)–A common fern from the Lower Cretaceous Crato Formation of northeast Brazil. Cretaceous Res. 2015; 54: 17–26. 76. Lindstro¨ m S, Irmis RB, Whiteside JH, Smith ND, Nesbitt SJ, Turner AH. Palynology of the upper Chinle Formation in northern New Mexico, U.S.A.: Implications for biostratigraphy and terrestrial ecosystem change during the Late Triassic (Norian–Rhaetian). Rev Palaeobot Palynol. 2016; 225: 106–131. 77. Thompson PW, Pflug H. Pollen und Sporen des Mitteleuropaïschen Tertia¨rs. Palaeontographica Abt B. 1953; 94 (1–4): 1–138. PLOS ONE

Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 44 / 51

78. Van Konijnenburg-Van Cittert JHA. In situ gymnosperm pollen from the Middle Jurassic of Yorkshire. Acta Bot Neerl. 1971; 20(1): 1–97. 79. Aze´ ma C, Boltenhagen E. Pollen du Cre´tace´ Moyen du Gabon attribue´ aux Ephedrales. Pale´ obiol Cont. 1974; 5(1): 1–37. 80. Alvin KL. Cheirolepidiaceae: Biology, structure and paleoecology. Rev Palaeobot Palynol. 1982; 37: 71–98. 81. Pedersen KR, Crane PR, Friis EM. Morphology and phylogenetic significance of Vardekloeftia Harris (Bennettitales). Rev Palaeobot Palynol. 1989; 60: 7–24. 82. Balme BE. Fossil in situ spores and pollen grains: an annotated catalogue. Rev Palaeobot Palynol. 1995; 87: 81–323. 83. Uličny´ D, Kvaček J, Svobodova´ M, Sˇ piča´ kova´ L. High-frequency sea-level fluctuations and plant habi- tats in Cenomanian fluvial to estuarine succession: Pecı´nov quarry, Bohemia. Palaeogeogr Palaeocli- matol Palaeoecol. 1997; 136: 165–197. 84. Stuchlik L, Ziembińska-Tworzydło M, Kohlman-Adamska A, Grabowska I, Ważyńska H, Sadowska A. Atlas of pollen and spores of the Polish Neogene. Volume 2 –Gymnosperms. Krako´ w: Polish Acad Sci; 2002. 85. Dejax J, Pons D, Yans J. Palynology of the dinosaur-bearing Wealden facies in the natural pit of Ber- nissart (Belgium). Rev Palaeobot Palynol. 2007; 144: 25–38. 86. Peyrot D, Barro´ n E, Comas-Rengifo MJ, Thouand E, Tafforeau P. A confocal laser scanning and con- ventional wide field light microscopy study of Circumpolles from the Toarcian-Aalenian of the Fuentel- saz section (Spain). Grana. 2007; 46: 217–226.

87. Peñalver E, Arillo A, Pe´rez de la Fuente R, Riccio M, Delclòs X, Barro´ n E, et al. Long-proboscid flies as pollinators of Cretaceous gymnosperms. Curr Biol. 2015; 25 (14): 1917–1923. https://doi.org/10. 1016/j.cub.2015.05.062 PMID: 26166781 88. Pattemore GA, Rigby JF, Playford G. Triassic-Jurassic pteridosperms of Australasia: speciation, diver- sity and decline. Bol Geol Min. 2015; 126 (4): 689–722. 89. Peris D, Pe´ rez de la Fuente R, Peñalver E, Delclòs X, Barro´ n E, Labandeira CC. False blister beetles and the expansion of gymnosperm-insect pollination modes before angiosperm dominance. Curr Biol. 2017; 27 (6): 897–904. https://doi.org/10.1016/j.cub.2017.02.009 PMID: 28262492 90. Kvaček J, Barro´n E, Heřmanova´ Z, Mendes MM, Karch J, Zˇ emlička J, et al. Araucarian conifer from the late Albian amber of northern Spain. Pap Palaeont. 2018; 4(4): 643–656. 91. Peyrot D, Barro´ n E, Polette F, Batten DJ, Ne´ raudeau D. Early Cenomanian palynofloras and inferred resiniferous forests and vegetation types in Charentes (southwestern France). Cretaceous Res. 2019; 94: 168–189. 92. Pacltova´ B. Some new pollen grains from the Bohemian Cenomanian. Rev Palaeobot Palynol. 1968; 7: 99–106. 93. Ward JV. Early Cretaceous angiosperm pollen from the Cheyenne and Kiowa Formations (Albian) of Kansas, U.S.A. Palaeontographica Abt B. 1986; 202 (1–6): 1–81. 94. Doyle JA, Endress PK, Upchurch GR. Early Cretaceous monocots: a phylogenetic evaluation. Acta Mus Natl Pragae B–Hist Nat. 2008; 64 (2–4): 59–87.

95. Raine JI, Mildenhall DC, Kennedy EM. New Zealand fossil spores and pollen: an illustrated catalogue. 4th edition. GNS Sci Misc Ser. 2011; 4. http://data.gns.cri.nz/sporepollen/index.htm 96. Stuchlik L, Ziembińska-Tworzydło M, Kohlman-Adamska A, Grabowska I, Słodkowska B, Worobiec E, et al. Atlas of pollen and spores of the Polish Neogene. Volume 4 –Angiosperms (2). Krako´ w: Polish Acad Sci; 2014. 97. Doyle JA, Endress PK. Integrating Early Cretaceous fossils into the phylogeny of living angiosperms: ANITA lines and relatives of Chloranthaceae. Int J Plant Sci. 2014; 175: 555–600. 98. Kvaček J, Doyle JA, Endress PK, Daviero-Gomez V, Gomez B, Tekleva M. Pseudoasterophyllites cre- taceous from the Cenomanian (Cretaceous) of the Czech Republic: A possible link between Chlor- anthaceae and Ceratophyllum. Taxon. 2016; 65(6): 1345–1373. 99. Llorens M, Pe´ rez Loinaze VS. Late Aptian angiosperm pollen grains from Patagonia: Earliest steps in flowering plant evolution at middle latitudes in southern South America. Cretaceous Res. 2016; 57: 66–78. 100. Lê S, Josse J, Husson F. (2008). FactoMineR: An R Package for Multivariate Analysis. J Stat Soft. 2008; 25 (1): 1–18. 101. Juggins S. Rioja: Analysis of Quaternary Science Data. R package version 0.9–26, Newcastle: New- castle Univ; 2020. PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 45 / 51

102. Corral JC, Lo´ pez Del Valle R, Alonso J. El a´ mbar creta´ cico de A´ lava (Cuenca Vasco-Canta´ brica, norte de España). Su colecta y preparacio´n. Estud Mus Cien Nat A´ lava. 1999; 14: 7–21. 103. Batten DJ. Chapter 26A. Palynofacies and Palaeoenvironmental interpretation. In: Jansonius J, McGregor DC, editors. Palynology: Principles and Applications, vol. 3. Salt Lake City: Am Assoc Stra- tigr Palynol Found; 1996. pp. 1011–1064. 104. Barro´ n E, Peyrot D, Rodrı´guez-Lo´ pez JP, Mele´ndez N, Lo´ pez del Valle R, Najarro M, et al. Palynology of Aptian and upper Albian (Lower Cretaceous) amber-bearing outcrops of the southern margin of the Basque-Cantabrian basin (northern Spain). Cretaceous Res. 2015; 52: 292–312. 105. Horikx M, Hochuli PA, Feist-Burkhardt S, Heimhofer U. Albian angiosperm pollen from shallow marine strata in the Lusitanian Basin, Portugal. Rev Palaeobot Palynol. 2016; 228: 67–92. 106. Gomez B, Martı´n-Closas C, Barale G, Sole´ de Porta N, The´venard F, Guignard G. Frenelopsis (Coni- ferales: Cheirolepidiaceae) and related male organ genera from the Lower Cretaceous of Spain. Palaeontology. 2002; 45(5): 997–1036. 107. Behrensmeyer AK, Hook RW. Paleoenvironmental contexts and taphonomic modes in the terrestrial fossil record, in: Behrensmeyer AK, Damuth JD, DiMichele WA, Potts R, Sues H-D, Wing SL, editors. Terrestrial Ecosystems through Time. Evolutionary paleoecology of terrestrial plants and animals. Chicago: Univ Chicago Press; 1992. pp. 15–38.

108. Peñalver E, Grimaldi DA, Delclòs X. Early Cretaceous spider web with its prey. Science. 2006; 312: 1761–1761. https://doi.org/10.1126/science.1126628 PMID: 16794072 109. Delclòs X, Peñalver E, Arillo A, Engel MS, Nel A, Azar D, et al. 2016. New mantises (Insecta: Manto- dea) in Cretaceous ambers from Lebanon, Spain, and Myanmar. Cretaceous Res. 2016; 60: 91–108. 110. Peñalver E, Ortega-Blanco J, Nel A, Delclòs X. Mesozoic Evaniidae (Insecta: Hymenoptera) in Span- ish amber: reanalysis of the phylogeny of the Evanioidea. Acta Geol Sin. 2010; 84 (4): 809–827. 111. Singh C. Cenomanian microfloras of the Peace River area, northwestern Alberta. Res Counc Alberta Bull. 1983; 44: 1–239. 112. Hochuli PA, Heimhofer U, Weissert H. Timing of early angiosperm radiation: recalibrating the classical succession. J Geol Soc Lond. 2006; 163: 587–594. 113. Hasenboehler B. E´ tude pale´obotanique et palynologique de l’Albien et du Cenomanien du "Bassin Occidental Portugais" au Sud de l’Accident de Nazare (Province d’Estremadure, Portugal). PhD The- sis. Universite´ Pierre et Marie Curie. 1981. 114. Bint AN. Fossil Ceratiaceae: A restudy and new taxa from the mid-Cretaceous of the Western Interior, U.S.A. Palynology. 1986; 10: 135–180. 115. Berthou PY, Leereveld H. Stratigraphic implications of palynological studies on Berriasian to Albian deposits from western and southern Portugal. Rev Palaeobot Palynol. 1990; 66: 313–344. 116. Schrank E, Ibrahim MIA. Cretaceous (Aptian–Maastrichtian) palynology of Foraminifera-dated wells

(KRM-1, AG-18) in northwestern Egypt. Berl Geowiss Abh, Reihe A. 1995; 177: 1–44. 117. Leereveld H, de Haan PJ, Juha´ sz M. Stratigraphic evaluation of spore/pollen assemblages from the Lower Cretaceous of the Alpine-Mediterranean Realm. Lab Palaeobot Palynol Contrib Ser. 1989; 89/ 07: 1–253+1–98. 118. Dea´ k MH, Combaz A. “Microfossiles organiques" du Wealdien et du Ce´nomanien dans un sondage de Charente-Maritime. Rev Micropale´ontol. 1967; 10(2): 69–96. 119. Ravn RL. Miospores from the Muddy Sandstone (upper Albian), Wind River Basin, Wyoming, U.S.A. Palaeontographica Abt B. 1995; 234: 41–91. 120. Pacltova´ B. Palynological study of Angiospermae from the Peruc Formation (?Albian-Lower Cenoma- nian) of Bohemia. Sb Geol Paleontol, řada P. 1971; 13: 105–139. 121. May FE. A survey of palynomorphs from several coal-bearing horizons of Utah. In: Doelling H.H., edi- tor. Utah Geol Min Surv Monogr Ser. 1972; 3: 497–542. 122. Ravn RL, Witzke BJ. The palynostratigraphy of the Dakota Formation (?late Albian–Cenomanian) in its type area, northwestern Iowa and northeastern Nebraska, USA. Palaeontographica Abt B. 1995; 234(3–6): 93–171. 123. Boulouard C, Cane´ rot J. Donne´ es nouvelles sur l’Aptien supe´ rieur et l’Albien dans le Bas-Arago´ n et le Maestrazgo (Espagne). Bull Cent Rech Pau-SNPA. 1970; 4(2): 453–463. 124. Sole´ de Porta N, Salas R. Conjuntos microflorı´sticos del Creta´ cico Inferior de la Cuenca del Maes- trazgo. Cordillera Ibe´rica Oriental (NE de España). Cuad Geol Ibe´ r. 1994; 18: 355–368.

125. Haq B. Cretaceous eustasy revisited. Global Planet Change. 2014; 113: 44–58. 126. Ju¨ rgens N. Floristic biodiversity and history of African arid regions. Biodiversity Conserv. 1997; 6 (3): 495–514. PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 46 / 51

127. Cowling RM, Rundel PW, Desmet PG, Esler KJ. Extraordinary high regional-scale plant diversity in southern Africa arid lands: subcontinental and global comparisons. Divers Distrib. 1998; 4: 27–36. 128. Muller J. Palynology of recent Orinoco delta and shelf sediments: reports of the Orinoco Shelf Expedi- tion; Volume 5. Micropaleontology. 1959; 5(1): 1–32. 129. Krasilov VA. Paleoecology of terrestrial plants. Basic principles and techniques. New York: John Wiley & sons, INC; 1975. 130. Spicer RA. Plant taphonomic processes. In: Allison PA, Briggs DE, editors. Taphonomy releasing: the data locked in the fossil record. New York: Plenum Press; 1991. pp. 71–113. 131. Davis MB. Palynology after Y2K - Understanding the source area of pollen in sediments. Annu Rev Earth Planet Sci. 2000; 28: 1–18. 132. Najarro M, Peñalver E, Pe´rez-de la Fuente R, Ortega-Blanco J, Menor-Salva´ n C, Barro´n E, et al. Review of the El Solplao amber outcrop, Early Cretaceous of Cantabria, Spain. Acta Geol Sin. 2010; 84 (4): 959–976. 133. Palynology Kimyai A. and biostratigraphy of the Lower Cretaceous sediments in the south barrow test well no. 1, point barrow, Alaska. Palynology. 2000; 24 (1): 201–215. 134. Tosolini A, McLoughlin S, Wagstaff B, Cantrill D, Gallagher S. Cheirolepidiacean foliage and pollen from Cretaceous high-latitudes of southeastern Australia. Gondwana Res. 2015; 27: 960–977. 135. Boukhamsin H, Peyrot D, Lang S, Vecoli M. Low-latitude? upper Barremian-lower Aptian palynoflora

and paleovegetation of the Biyadh Formation (Arabian Plate, eastern margin of northern Gondwana): evidence for a possible cold nap. Cretaceous Res. 2022a; 129: 104995. 136. Peyrot D, Playford G, Mantle DJ, Backhouse J, Milne LA, Carpenter RJ, et al. The greening of Western Australian landscapes: the Phanerozoic plant record. J R Soc West Aust. 2019; 102: 52–82. 137. Boltenhagen E., Salard-Cheboldaeff M., 1987. E´ tude palynologique du sel aptien du Congo. Me´m Trav Inst Montpellier Ecole Pratique Hautes Etud. 1987; 17: 273–293. 138. Watson J. Some Lower Cretaceous conifers of the Cheirolepidiaceae from the U.S.A. and England. Palaeontology. 1977; 20: 715–749. 139. Watson J. The Cheirolepidiaceae. In: Beck CB, editor. Origin and evolution of gymnosperms. New York: Columbia Univ Press; 1988. pp. 382–447. 140. Hotton CL, Baghai-Riding NL. Palynological evidence for conifer dominance within a heterogeneous landscape in the Late Jurassic Morrison Formation, U.S.A. In: Gee CT, editor. Plants in Mesozoic time: Morphological innovations, phylogeny, ecosystems. Bloomington: Indiana Univ Press; 2010. pp. 295–328. 141. Francis JE. The dominant conifer of the Jurassic Purbeck Formation, England. Palaeontology. 1983; 26: 277–294. 142. McLoughlin S, Tosolini A-MP, Nagalingum NS, Drinnan AN. Early Cretaceous (Neocomian) flora and fauna of the lower Strzelecki Group, Gippsland Basin, Victoria. Mem Assoc Australasian Palaeontol. 2002; 26: 1–144. 143. Farjon A. A monograph of Cupressaceae and Sciadopitys. Kew: Royal Botanic Gardens; 2005.

144. Pelzer G, Riegel W, Wilde V. Depositional controls on the Lower Cretaceous Wealden coals of north- west Germany. In: McCabe PJ, Parrish JT, editors. Controls on the Distribution and Quality of Creta- ceous Coals. Geol Soc Am Spec Pap. 1992; 267: 227–243. 145. Mays C. A Late Cretaceous (Cenomanian–Turonian) south polar palynoflora from the Chatham Islands, New Zealand. Mem Assoc Australas Palaeontol. 2015; 47: 1–92. 146. McIver EE. Cretaceous Widdringtonia Endl. (Cupressaceae) from North America. Int J Plant Sci. 2001; 162 (4): 937–961. 147. Hamad A, Abu M, Amireh B, Jasper A, Uhl D. New palaeobotanical data from the Jarash Formation (Aptian–Albian, Kurnub Group) of NW Jordan. The Palaeobotanist. 2016; 65: 19–29. 148. Hamad AMA, Amireh B, El Atfy H, Jasper A, Uhl D. Fire in a Weichselia-dominated coastal ecosystem from the Lower Cretaceous (Barremian) of the Kurnub Group in NW Jordan. Cretaceous Res. 2016; 66: 82–93. 149. Dettmann ME. Cretaceous vegetation: the microfossil record. In: Hill RS, editor. History of the Austra- lian vegetation: Cretaceous to recent. Adelaide: Univ Adelaide Press; 2017. pp. 143–170. 150. Blanco-Moreno C, Gomez B, Buscalioni A´ . Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia. Geobios. 2018; 51: 571–578. 151. Halamski AT, Kvaček J, Svobodova´ M, Durska E, Heřmanova´ Z. Late Cretaceous mega-, meso-, and microfloras from Lower Silesia. Acta Palaeontol Pol. 2020; 65(4): 811–878. PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 47 / 51

152. Feild TS, Arens NC, Doyle JA, Dawson TE, Donoghue MJ. Dark and disturbed: a new image of early angiosperm ecology. Paleobiology. 2004; 30: 82–107. 153. Gomez B, Coiffard C, Sender LM, Martı´n-Closas C, Villanueva-Amadoz U, Ferrer J. Klitzschophyllites, aquatic basal eudicots (Ranunculales?) from the upper Albian (Lower Cretaceous) of northeastern Spain. Int J Plant Sci. 2009; 170 (8): 1075–1085. 154. Doyle JA, Endress PK. Integrating Early Cretaceous fossils into the phylogeny of living angiosperms: Magnoliidae and eudicots. J Syst Evol. 2010; 48(1): 1–35. 155. Feild T, Chatelet D, Brodribb T. Ancestral xerophobia: a hypothesis on the whole plant ecophysiology of early angiosperms. Geobiology. 2009; 7: 237–264. https://doi.org/10.1111/j.1472-4669.2009. 00189.x PMID: 19260972 156. Taylor DW, Hu S. Coevolution of early angiosperms and their pollinators: Evidence for pollen. Palaeontographica Abt B. 2010; 283: 103–135. 157. Vidiella PE, Armesto JJ, Gutie´ rrez JR. Vegetation changes and sequential flowering after rain in the southern Atacama Desert. J Arid Environ. 1999; 43: 449–458. 158. Elliot MB. Modern pollen–vegetation relationships in Northland, New Zealand. N Z J Bot. 1999; 37: 131–148. 159. Sousa VA, Hattemer HH. Pollen dispersal and gene flow by pollen in Araucaria angustifolia. Aust J Bot. 2003; 51: 309–317. 160. Chaler R, Grimalt JO. Fingerprint of Cretaceous higher plant resins by inferred spectroscopy and gas chromatography coupled to mass spectrometry. Phytochem Anal. 2005; 16(6): 446–450.

161. Dal Corso J, Roghi G, Ragazzi E, Angelini I, Giaretta A, Soriano C, et al. Physico-chemical analysis of Albian (Lower Cretaceous) amber from San Just (Spain): implications for palaeoenvironmental and palaeoecological studies. Geol Acta. 2013; 11(3): 359–370. 162. Peris D, Davis SR, Engel MS, Delclòs X. An evolutionary history embedded in amber: reflection of the Mesozoic shift in weevil-dominated (Coleoptera: Curculionoidea) faunas. Zool J Linn Soc. 2014; 171: 534–553. 163. Distribution Jaffre´ T. and ecology of conifers in New Caledonia. In: Enright NJ, Hill RS, editors. Distri- bution and ecology of southern conifers. Carlton: Melbourne Univ Press; 1995. pp. 171–196. 164. Ogden J, Stewart GH. Community dynamics of the New Zealand conifers. In: Enright NJ, Hill RS, edi- tors. Distribution and ecology of southern conifers. Carlton: Melbourne Univ Press; 1995. pp. 81–119. 165. Farjon A. A natural history of conifers. London: Timber Press; 2008. 166. Jackson ST, Overpeck JT. Responses of plant population and communities to environmental changes of the late Quaternary. Paleobiology. 2000; 26 (4): 194–220. 167. Time Pillon Y. and tempo of diversification in the flora of New Caledonia. Bot J Linn Soc. 2012; 170: 288–298. 168. Arillo A, Peñalver E, Delclòs X. Microphorites (Diptera: Dolichopodidae) from the Lower Cretaceous amber of San Just (Spain), and the co-occurrence of two ceratopogonid species in Spanish amber deposits. Zootaxa. 2008; 1920: 29–40. 169. Nel A., Perrichot V., Daugeron C., Ne´raudeau D. 2004. A new Microphorites in the Lower Cretaceous

amber of the southwest of France (Diptera: Dolichopodidae, “Microphorinae”). Ann Soc Entomol Fr. 2004; 40 (1): 23–29. 170. Ecology Regal P. and the evolution of flowering plant dominance. Science. 1977; 1996: 622–629. 171. Upchurch GR Jr, Doyle JA. Paleoecology of the conifers Frenelopsis and Pseudofrenelopsis (Cheiro- lepidiaceae) from the Cretaceous Potomac Group of Maryland and Virginia. In: Romans RC, editor. Geobotany II. New York: Plenum Press; 1981. pp. 167–202. 172. Kvaček J. Frenelopsis alata and its microsporangiate and ovuliferous reproductive structures from the Cenomanian of bohemia (Czech Republic, Central Europe). Rev Palaeobot Palynol. 2000; 112: 51– 78. https://doi.org/10.1016/s0034-6667(00)00035-x PMID: 11042326 173. Daviero V, Gomez B, Philippe M. Uncommon branching pattern within conifers: Frenelopsis turolensis, a Spanish Early Cretaceous Cheirolepidiaceae. Can J Bot. 2001; 79: 1400–1408. 174. McElwain JC, Chaloner WG. The Fossil Cuticle as a Skeletal Record of Environmental Change. Palaios. 1996; 11(4): 376–388. 175. Haworth M, McElwain J. Hot, dry, wet, cold or toxic? Revisiting the ecological significance of leaf and cuticular micromorphology. Palaeogeogr Palaeoclimatol Palaeoecol. 2008; 262(1): 79–90. 176. Gomez B, Coiffard C, De´ pre´ E´ , Daviero-Gomez V, Ne´ raudeau D. Diversity and histology of a plant lit- ter bed from the Cenomanian of Archingeay-Les Nouillers (southwestern France). CR Palevol. 2008; 7: 135–144. PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 48 / 51

177. Hlusˇtı´k A. Eretmophyllous Ginkgoales from the Cenomanian. Acta Mus Natl Pragae, ser B Hist Nat. 1986; 42(1–2): 99–115. 178. Kvaček J, Falcon-Lang HJ, Dasˇkova´ J. A new Late Cretaceous ginkgoalean reproductive structure Nehvizdyella gen. nov. from the Czech Republic and its whole-plant reconstruction. Am J Bot. 2005; 92(12): 1958–1969. 179. Hughes RG. Theories and models of species abundance. Am Nat. 1986; 128: 879–899. 180. Kvaček J, Gomez B, Zetter R. The early angiosperm Pseudoasterophyllites cretaceus from Albian- Cenomanian of the Czech Republic and France revisited. Acta Palaeontol Pol. 2012; 57(2): 437–443. 181. Kvaček J. Late Cretaceous floras in Central Europe and their palaeoenvironment. Praha: Univ Karlovy v Praze; 2017. 182. Krassilov VA. On Montsechia, an angiospermoid plant from Lower Cretaceous of Las Hoyas, Spain: new data and interpretations. Acta Palaeobot. 2011; 51(2): 181–205. 183. Friis EM, Pedersen K, Crane P. Cretaceous diversification of angiosperms in the western part of the Iberian Peninsula. Rev Palaeobot Palynol. 2010; 162: 341–361. 184. Heimhofer U, Hochuli PA, Burla S, Weissert H. New records of Early Cretaceous angiosperm pollen from Portuguese coastal deposits: Implications for the timing of the early angiosperm radiation. Rev Palaeobot Palynol. 2007; 144: 39–76. 185. Hughes N. The Enigma of Angiosperms origins. Cambridge: Cambridge Univ Press; 1994. 186. Regali M., Tucanopollis a new genus of early angiosperms. Bol Geoci Petrobra´s. 1989; 3: 395–402.

187. Doyle J. Revised palynological correlations of the lower Potomac Group (USA) and the Cocobeach sequence of Gabon (Barremian–Aptian). Cretaceous Res. 1992; 13: 337–349. 188. Rodrı´guez-Lo´pez JP, Mele´ ndez N, De Boer PL, Soria AR. Controls on marine–erg margin cycle vari- ability: aeolian–marine interaction in the mid-Cretaceous Iberian Desert System, Spain. Sedimentol- ogy. 2012; 59(2): 466–501. 189. Rodrı´guez-Lo´pez JP, Barro´ n E, Peyrot D, Hughes GB. Deadly oasis: Recurrent annihilation of Creta- ceous desert bryophyte colonies; the role of solar, climate and lithospheric forcing. Geosci Front. 2021; 12: 1–12. 190. Engel MS, Delclòs X. Primitive termites in Cretaceous amber from Spain and Canada (Isoptera). J Kansas Entomol Soc. 2010; 83 (2): 111–128. 191. Peris D, Philips TK, Delclòs X. Ptinid beetles from the Cretaceous gymnosperm-dominated forests. Cretaceous Res. 2015; 52(Part B): 440–452. 192. Arillo A, Subı´as LS, Sa´nchez-Garcı´a A. New species of fossil oribatid mites (Acariformes, Oribatida), from the Lower Cretaceous amber of Spain. Cretaceous Res. 2016; 63: 68–76. 193. Retallack G J. Permian–Triassic life crisis on land. Science. 1995; 267 (5194): 77–80. https://doi.org/ 10.1126/science.267.5194.77 PMID: 17840061 194. Vajda V, Raine JI, Hollis CJ. Indication of global deforestation at the Cretaceous–Tertiary boundary by New Zealand fern spike. Science. 2001; 294 (5547): 1700–1702. https://doi.org/10.1126/science. 1064706 PMID: 11721051 195. Van de Schootbrugge B, Quan TM, Lindstrom SL, Pu¨ ttmann W, Heunisch C, Pross J, et al. Floral

changes across the Triassic/Jurassic boundary linked to flood basalt volcanism. Nat Geosci. 2009; 2 (8): 589–594. 196. Spicer RA. Changing climate and biota. In: Skelton PW, editor. The Cretaceous World. Cambridge: Cambridge Univ Press; 2003. pp. 85–162. 197. Brown JH, Valone TJ, Curtin CG 1997. Reorganization of an arid ecosystem in response to recent cli- mate change. PNAS. 1997; 94 (18): 9729–9733. https://doi.org/10.1073/pnas.94.18.9729 PMID: 11038570 198. Van de Koppel J, Rietkerk M. Spatial Interactions and Resilience in Arid Ecosystems. Am Nat. 2004; 163(1): 113–121. https://doi.org/10.1086/380571 PMID: 14767841 199. Peyrot D, Rodrı´guez-Lo´pez JP, Lassaletta L, Mele´ ndez N, Barro´ n E. Contributions to the palaeoenvir- onmental knowledge of the Escucha Formation in the Lower Cretaceous Oliete Sub-basin, Teruel, Spain. CR Palevol. 2007; 6: 469–481. 200. Page CN. Ecological strategies in fern evolution: a neopteridological overview. Rev Palaeobot Palynol. 2002; 119: 1–33. 201. Hietz P. Fern adaptations to xeric environments. In: Mehltreter K, Walker L, Sharpe J, editors. Fern Ecology. Cambridge: Cambridge Univ Press; 2012. pp. 140–176. PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 49 / 51

202. Vitt DH, Crandall-Stotler B, Wood AJ. Bryophytes, survival in a dry world through tolerance and avoid- ance. In: Rajakaruna N, Boyd RS, Harris TB, editors. Plant Ecology and Evolution in Harsh Environ- ments. New York: Nova Science; 2014. pp. 267–295. 203. Doyle J, Endress PK. Phylogenetic analyses of Cretaceous fossils related to Chloranthaceae and their evolutionary implications. Bot Rev. 2018; 84: 156–202. 204. Coiffard C, Gomez B, Thie´ baut M, Kvaček J, The´venard F, Ne´ raudeau D. Inframarginal veined Laura- ceae leaves from the Albian–Cenomanian of Charente-Maritime (western France). Palaeontology. 2009; 52(2): 323–336. 205. Crabtree DR. Mid-Cretaceous ferns in situ from the Albino member of the Mowry Shale, southwestern Montana. Palaeontographica Abt. B 1988; 209: 1–27. 206. Ahlbrandt TS, Fryberger SG. Sedimentary features and significance of interdune deposits. Soc Econ Paleontol Min Spec Publ. 1981; 31: 293–314. 207. Mudie PJ, McCarthy FMG. Late Quaternary pollen transport processes, western North Atlantic: Data from box models, cross-margin and N-S transects. Mar Geol. 1994; 118: 79–105. 208. Mudie PJ, McCarthy FMG. Marine palynology: potentials for onshore-offshore correlation of Pleisto- cene–Holocene records. Trans R Soc South Afr. 2006; 61: 139–157. 209. Chaloner WG, Muir M. Spores and Floras. In: Marendson L, Westord J, editors. Coal and coal-bearing strata. Edinburgh: Oliver and Boyd; 1968. pp. 127–146. 210. Ibrahim MIA. Aptian–Turonian palynology of the Ghazalat-1 Well (GTX-1) Qattara Depression, Egypt.

Rev Palaeobot Palynol. 1996; 94: 137–168. 211. Ibrahim MIA. Late Albian–middle Cenomanian palynofacies and palynostratigraphy, Abu Gharadig-5 well, Western Desert, Egypt. Cretaceous Res. 2002; 2: 775–788. 212. El-Beialy S, El-Soughier M, Mohsen SA, El Atfy H. Palynostratigraphy and paleoenvironmental signifi- cance of the Cretaceous succession in the Gebel Rissu-1 well, north Western Desert, Egypt. J Afr Earth Sci. 2011; 59: 215–226. 213. Schrank E. Palaeoecological aspects of Afropollis/elaterates peaks (Albian–Cenomanian pollen) in the Cretaceous of northern Sudan and Egypt. In: Goodman DK, Clarke RT, editors. Proc IX Int Palynol Congress, Houston, Texas, USA, 1996. Am Assoc Stratigr Palynol Found; 2001. pp. 201–210. 214. Schrank E. Palynology of the Albian Makhtesh Qatan site, northern Negev (Israel), with descriptions of two new pollen species. Rev Palaeobot Palynol. 2017; 246: 185–215. 215. Shih PJ, Li L, Li D, Ren D. Application of geometric morphometric analyses to confirm three new wasps of Evaniidae (Hymenoptera: Evanioidea) from mid-Cretaceous Myanmar amber. Cretaceous Res. 2020; 109: 104249. 216. Engel MS, Perrichot V. The extinct wasp family Serphitidae in late Cretaceous Vendean amber (Hymenoptera). Paleontol Contrib. 2014; 10J: 46–51. 217. Me´ dus J, Berthou PY. Palynoflores dans la coupe de l’Albien de Foz do Folcão (Portugal). Geobios. 1980; 13: 263–269. 218. Pons D, Vozenin-Serra C. Wood of Ginkgoales in the Cenomanian of Anjou, France. Cour Forchinst Senckenb. 1992; 147: 199–213.

219. Gomez B, Daviero-Gomez V, Perrichot V, The´ venard F, Coiffard C, Philippe M, et al. Assemblages floristiques de l’Albien-Ce´nomanien de Charente-Maritime (SO France). Ann Paleontol. 2004; 90: 147–159. 220. Ne´ raudeau D, Vullo R, Gomez B, Perrichot V, Videt B. Stratigraphie et pale´ontologie (plantes, verte´ - bre´s) de la se´ rie paralique Albien terminal-Ce´nomanian basal de Tonnay-Charente (Charente-Mari- time, France). CR Palevol. 2005; 4: 79–93. 221. Spicer B, Skelton PW. The operation of the major geological carbon sinks. In: Skelton PW, editor. The Cretaceous World. Cambridge: Univ Press; 2003. pp. 249–271. 222. Brenner GJ. The spores and pollen of the Potomac Group of Maryland. Maryland Dep Geol Mines Water Res Bull. 1963; 27: 1–215. 223. Kemp EM. Aptian and Albian miospores from southern England. Palaeontographica Abt. B 1970; 131 (1–4): 73–143. 224. Svobodova´ M. Mid-Cretaceous palynomorphs from the Blansko Graben (Czech Republic): affinities to both Tethyan and Boreal bioprovinces. NR 58 Proc 4th EPPC: Mid-Cretaceous palynomorphs from the Blansko Graben; 1997. pp. 149–155. 225. Svobodova´ M, Hradecka´ L, Skupien P, Sˇ va´ benicka´ L. Microfossils of theAlbian and Cenomanian shales from the Stramberk area (Silesia unit, Outer Western Carpathians, Czech Republic). Geol Car- path. 2004; 55 (5): 371–388. PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 50 / 51

226. Polette F, Licht A, Cincotta A, Batten DJ, Depuydt P, Ne´ raudeau D, et al. Palynological assemblage from the lower Cenomanian plant-bearing Lagersta¨ tte of Jaunay-Clan-Ormeau-Saint-Denis (Vienne, western France): Stratigraphic and paleoenvironmental implications. Rev Palaeobot Palynol. 2019; 271: 104102. 227. Spores Norris G. and pollen from the Lower Colorado Group (?Albian–Cenomanian) of central Alberta. Palaeontographica Abt B. 1967; 120 (1–4): 72–115. 228. Playford G. Palynology of Lower Cretaceous (Swan River) strata of Saskatchewan and Manitova. Palaeontology. 1971; 14 (4): 533–565. 229. Singh C. Lower Cretaceous microfloras of the Peace River Area, northwestern Alberta. Res Counc Alberta Bull. 1971; 28 (1): 1. PLOS ONE Late Albian desert biota from Iberia PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 51 / 51