Ari Iglesias
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The organizations/institutes from the scholar's latest article
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Research Interests
Ari Iglesias has a total of 161 co-authors. Here are the top co-authors by number of shared publications:
- Peter Wilf
Department of Geosciences and Earth and Environmental Systems Institute Pennsylvania State University University Park PA 16802 USA
10 shared publications N. Rubén Cúneo
CONICET-Museo Paleontológico Egidio Feruglio, Av. Fontana 140, 9100, Trelew, Chubut, Argentina
7 shared publicationsPeter Wilf
Department of Geosciences, Pennsylvania State University, University Park, PA, 16802, USA
6 shared publications- Daniel Gustavo Poiré
Universidad Nacional de La Plata, Facultad de Ciencias Naturales y Museo, Centro de Investigaciones Geológicas, La Plata, Argentina
5 shared publications Kirk R. Johnson
4 shared publicationsAlba B. Zamuner
3 shared publications
Latest Publications
- Open Access
- 9 September 2026
- Published by Wiley in American Journal of Botany
- Vol. 113 (9) , e70253
Austrocedrus from the Late Cretaceous of Antarctica and its implications for the early evolution of southern cypresses (Callitroideae, Cupressaceae)
Premise: The modern distribution of the southern cypresses, Callitroideae (Cupressaceae), reflects a classic Gondwanan distribution. The initial diversification of Callitroideae is estimated to have occurred during the Cretaceous, yet fossils from this period are rare, hampering our understanding of its early evolutionary history. Methods: A permineralized cluster of callitroid seed cones has been recovered from the Late Cretaceous Santa Marta Formation on James Ross Island, Antarctica. The specimen was sectioned using the acetate peel technique. We conducted a comparative morphological and anatomical investigation of the fossil with living members of Callitroideae and tested its phylogenetic placement using Bayesian and parsimony inference methods. Results: Each cone has four opposite‐decussate ovuliferous complexes, a central columella, and subtending bifacially flattened shoots. This new fossil resembles living Austrocedrus in having ovuliferous complexes with a short, apically located bract tip and an abaxial row of vascular bundles. Phylogenetic reconstructions place the fossil sister to Austrocedrus chilensis , and we erect a new species, Austrocedrus vetus sp. nov. Conclusions: Austrocedrus is known by a handful of Cenozoic fossils from Australia and a single surviving taxon, A. chilensis , in the Southern Andes of Chile and Argentina. Austrocedrus vetus is the oldest evidence of the genus, extending its stratigraphic record by 40 Ma. Our findings indicate that Austrocedrus diverged by the Campanian and experienced stasis in seed cone morphology. Furthermore, this fossil is the first occurrence of Austrocedrus in Antarctica, demonstrating that the genus was an element of Antarctic forest ecosystems during the Late Cretaceous.
- 9 September 2026
- Published by Wiley in American Journal of Botany
- Vol. 113 (9) , e70253
Open AccessAustrocedrus from the Late Cretaceous of Antarctica and its implications for the early evolution of southern cypresses (Callitroideae, Cupressaceae)
Premise: The modern distribution of the southern cypresses, Callitroideae (Cupressaceae), reflects a classic Gondwanan distribution. The initial diversification of Callitroideae is estimated to have occurred during the Cretaceous, yet fossils from this period are rare, hampering our understanding of its early evolutionary history. Methods: A permineralized cluster of callitroid seed cones has been recovered from the Late Cretaceous Santa Marta Formation on James Ross Island, Antarctica. The specimen was sectioned using the acetate peel technique. We conducted a comparative morphological and anatomical investigation of the fossil with living members of Callitroideae and tested its phylogenetic placement using Bayesian and parsimony inference methods. Results: Each cone has four opposite‐decussate ovuliferous complexes, a central columella, and subtending bifacially flattened shoots. This new fossil resembles living Austrocedrus in having ovuliferous complexes with a short, apically located bract tip and an abaxial row of vascular bundles. Phylogenetic reconstructions place the fossil sister to Austrocedrus chilensis , and we erect a new species, Austrocedrus vetus sp. nov. Conclusions: Austrocedrus is known by a handful of Cenozoic fossils from Australia and a single surviving taxon, A. chilensis , in the Southern Andes of Chile and Argentina. Austrocedrus vetus is the oldest evidence of the genus, extending its stratigraphic record by 40 Ma. Our findings indicate that Austrocedrus diverged by the Campanian and experienced stasis in seed cone morphology. Furthermore, this fossil is the first occurrence of Austrocedrus in Antarctica, demonstrating that the genus was an element of Antarctic forest ecosystems during the Late Cretaceous.
- Open Access
- 17 July 2026
- Published by Wiley in New Phytologist
- Vol. 251 (6) , 3627-3639
Cretaceous moss–cyanobacteria association reveals trophic interactions in polar forests of Antarctica
Summary: Through symbiotic associations with microbes, mosses play many important ecological roles in terrestrial ecosystems, such as facilitating trophic interactions and nitrogen cycling. However, little is known about interactions between mosses and microbes in past ecosystems. Here, we characterize a permineralized moss with epiphytic cyanobacteria and tubular hyphae from the Cretaceous of Antarctica. The fossil was identified in a calcium carbonate concretion from the Beta member of the Santa Marta Formation Cretaceous (Campanian) from James Ross Island. The fossil was studied using acetate peels and light microscopy. The fossil moss is assignable to Bryopsida. Cyanobacteria trichomes with heterocyst‐like cells are present in leaf axils and the perichaetium. In some leaf axils, epiphytic fungi are preserved. The apical perichaetium contains several archegonia; one contains an egg cell while another contains a possible embryo. Fossil epiphytic cyanobacteria demonstrate that moss associations were part of Antarctic polar forests during the Cretaceous. Recognizing their importance in modern ecosystems, moss–cyanobacteria associations likely contributed bioavailable nitrogen to the nutrient cycle of ancient polar forests, which would benefit organisms in different trophic levels. This work reveals long‐term stasis in commensal relationships between cyanobacteria and mosses and provides a minimum age for these relationships.
- 17 July 2026
- Published by Wiley in New Phytologist
- Vol. 251 (6) , 3627-3639
Open AccessCretaceous moss–cyanobacteria association reveals trophic interactions in polar forests of Antarctica
Summary: Through symbiotic associations with microbes, mosses play many important ecological roles in terrestrial ecosystems, such as facilitating trophic interactions and nitrogen cycling. However, little is known about interactions between mosses and microbes in past ecosystems. Here, we characterize a permineralized moss with epiphytic cyanobacteria and tubular hyphae from the Cretaceous of Antarctica. The fossil was identified in a calcium carbonate concretion from the Beta member of the Santa Marta Formation Cretaceous (Campanian) from James Ross Island. The fossil was studied using acetate peels and light microscopy. The fossil moss is assignable to Bryopsida. Cyanobacteria trichomes with heterocyst‐like cells are present in leaf axils and the perichaetium. In some leaf axils, epiphytic fungi are preserved. The apical perichaetium contains several archegonia; one contains an egg cell while another contains a possible embryo. Fossil epiphytic cyanobacteria demonstrate that moss associations were part of Antarctic polar forests during the Cretaceous. Recognizing their importance in modern ecosystems, moss–cyanobacteria associations likely contributed bioavailable nitrogen to the nutrient cycle of ancient polar forests, which would benefit organisms in different trophic levels. This work reveals long‐term stasis in commensal relationships between cyanobacteria and mosses and provides a minimum age for these relationships.
- 12 June 2026
- Published by Wiley in American Journal of Botany
- Vol. 113 (6) , e70220
Fossil fruits of a new Attaleinae (Cocoseae, Arecaceae): Evidence of a palm grove community during the Early Miocene in northern Patagonia, Argentina
Premise: Palms can develop a wide variety of fruit shapes and sizes. Species in the Cocoseae tribe possess an evident synapomorphy: the presence of three “eyes” or germination pores in their endocarp, probably best known by the widespread monotypic extant Cocos nucifera . There is little consensus on the biogeographic history of the cocosoid palms, there being few previous world‐spread fossil records, some of which require further study in order to establish a clear taxonomic assignment. Methods: Anatomy of permineralized fruits (pyrenes) from the Miocene in northern Patagonia, Argentina, was studied using thin sectioning and polished surfaces observed under transmitted and epifluorescence microscopy. Some fossils were analyzed under X‐ray micro‐tomography. Results: We report a new palm fruit fossil record based on anatomical studies of >30 specimens. These were assigned to a new fossil taxon within the subtribe Attaleinae (Tribe Cocoseae, Arecoideae). Furthermore, the presence of an unusual character as endocarp intrusions suggests a close anatomical relationship with the extant Syagrus romanzoffiana . Conclusions: The new fossil taxon is one of the best‐preserved fossils of the Cocoseae worldwide and is the first unambiguous fossil record for Attaleinae from southern South America. The extremely high concentration (hundreds to thousands) of pyrenes within several strata of the basal section of the Chichinales Formation and their wide distribution in the study area allow for the interpretation of a palm grove community. This record represents the most austral occurrence of the Attaleinae in South America, indicating that subtropical climatic conditions extended into northern Patagonia almost to the Early Miocene.
- 12 June 2026
- Published by Wiley in American Journal of Botany
- Vol. 113 (6) , e70220
Fossil fruits of a new Attaleinae (Cocoseae, Arecaceae): Evidence of a palm grove community during the Early Miocene in northern Patagonia, Argentina
Premise: Palms can develop a wide variety of fruit shapes and sizes. Species in the Cocoseae tribe possess an evident synapomorphy: the presence of three “eyes” or germination pores in their endocarp, probably best known by the widespread monotypic extant Cocos nucifera . There is little consensus on the biogeographic history of the cocosoid palms, there being few previous world‐spread fossil records, some of which require further study in order to establish a clear taxonomic assignment. Methods: Anatomy of permineralized fruits (pyrenes) from the Miocene in northern Patagonia, Argentina, was studied using thin sectioning and polished surfaces observed under transmitted and epifluorescence microscopy. Some fossils were analyzed under X‐ray micro‐tomography. Results: We report a new palm fruit fossil record based on anatomical studies of >30 specimens. These were assigned to a new fossil taxon within the subtribe Attaleinae (Tribe Cocoseae, Arecoideae). Furthermore, the presence of an unusual character as endocarp intrusions suggests a close anatomical relationship with the extant Syagrus romanzoffiana . Conclusions: The new fossil taxon is one of the best‐preserved fossils of the Cocoseae worldwide and is the first unambiguous fossil record for Attaleinae from southern South America. The extremely high concentration (hundreds to thousands) of pyrenes within several strata of the basal section of the Chichinales Formation and their wide distribution in the study area allow for the interpretation of a palm grove community. This record represents the most austral occurrence of the Attaleinae in South America, indicating that subtropical climatic conditions extended into northern Patagonia almost to the Early Miocene.
- Open Access
- 1 January 2024
- Published by Polska Akademia Nauk Instytut Paleobiologii (Institute of Paleobiology, Polish Academy of Sciences) in Acta Palaeontologica Polonica
- Vol. 69 (3) , 467-483
A Late Jurassic deep-bodied actinopterygian from Antarctica
- 1 January 2024
- Published by Polska Akademia Nauk Instytut Paleobiologii (Institute of Paleobiology, Polish Academy of Sciences) in Acta Palaeontologica Polonica
- Vol. 69 (3) , 467-483
Open AccessA Late Jurassic deep-bodied actinopterygian from Antarctica
- Open Access
- 2 May 2023
- Published by Wiley in American Journal of Botany
- Vol. 110 (5) , e16169
The first Gondwanan Euphorbiaceae fossils reset the biogeographic history of the Macaranga‐Mallotus clade
Premise: The spurge family Euphorbiaceae is prominent in tropical rainforests worldwide, particularly in Asia. There is little consensus on the biogeographic origins of the family or its principal lineages. No confirmed spurge macrofossils have come from Gondwana.Methods: We describe the first Gondwanan macrofossils of Euphorbiaceae, represented by two infructescences and associated peltate leaves from the early Eocene (52 Myr ago [Ma]) Laguna del Hunco site in Chubut, Argentina.Results: The infructescences are panicles bearing tiny, pedicellate, spineless capsular fruits with two locules, two axile lenticular seeds, and two unbranched, plumose stigmas. The fossils' character combination only occurs today in some species of the Macaranga‐Mallotus clade (MMC; Euphorbiaceae), a widespread Old‐World understory group often thought to have tropical Asian origins. The associated leaves are consistent with extant Macaranga.Conclusions: The new fossils are the oldest known for the MMC, demonstrating its Gondwanan history and marking its divergence by at least 52 Ma. This discovery makes an Asian origin of the MMC unlikely because immense oceanic distances separated Asia and South America 52 Ma. The only other MMC reproductive fossils so far known are also from the southern hemisphere (early Miocene, southern New Zealand), far from the Asian tropics. The MMC, along with many other Gondwanan survivors, most likely entered Asia during the Neogene Sahul‐Sunda collision. Our discovery adds to a substantial series of well‐dated, well‐preserved fossils from one undersampled region, Patagonia, that have changed our understanding of plant biogeographic history.
- 2 May 2023
- Published by Wiley in American Journal of Botany
- Vol. 110 (5) , e16169
Open AccessThe first Gondwanan Euphorbiaceae fossils reset the biogeographic history of the Macaranga‐Mallotus clade
Premise: The spurge family Euphorbiaceae is prominent in tropical rainforests worldwide, particularly in Asia. There is little consensus on the biogeographic origins of the family or its principal lineages. No confirmed spurge macrofossils have come from Gondwana.Methods: We describe the first Gondwanan macrofossils of Euphorbiaceae, represented by two infructescences and associated peltate leaves from the early Eocene (52 Myr ago [Ma]) Laguna del Hunco site in Chubut, Argentina.Results: The infructescences are panicles bearing tiny, pedicellate, spineless capsular fruits with two locules, two axile lenticular seeds, and two unbranched, plumose stigmas. The fossils' character combination only occurs today in some species of the Macaranga‐Mallotus clade (MMC; Euphorbiaceae), a widespread Old‐World understory group often thought to have tropical Asian origins. The associated leaves are consistent with extant Macaranga.Conclusions: The new fossils are the oldest known for the MMC, demonstrating its Gondwanan history and marking its divergence by at least 52 Ma. This discovery makes an Asian origin of the MMC unlikely because immense oceanic distances separated Asia and South America 52 Ma. The only other MMC reproductive fossils so far known are also from the southern hemisphere (early Miocene, southern New Zealand), far from the Asian tropics. The MMC, along with many other Gondwanan survivors, most likely entered Asia during the Neogene Sahul‐Sunda collision. Our discovery adds to a substantial series of well‐dated, well‐preserved fossils from one undersampled region, Patagonia, that have changed our understanding of plant biogeographic history.
- 14 October 2022
- Published by Elsevier in Cretaceous Research
- Vol. 142, 105395
The Valcheta Petrified Forest (Upper Cretaceous), northern Patagonia, Argentina: A geological and paleobotanical survey
- 14 October 2022
- Published by Elsevier in Cretaceous Research
- Vol. 142, 105395
The Valcheta Petrified Forest (Upper Cretaceous), northern Patagonia, Argentina: A geological and paleobotanical survey
- Open Access
- 1 January 2022
- Published by FapUNIFESP (SciELO) in Anais da Academia Brasileira de Ciências
- Vol. 94 (suppl 1) , e20211142
Late Campanian-Early Maastrichtian Vertebrates From The James Ross Basin, West Antarctica: Updated Synthesis, Biostratigraphy, And Paleobiogeography
The Snow Hill Island Formation (SHIF; late Campanian - early Maastrichtian) crops out in the northeast of the Antarctic Peninsula and constitutes the basal part of the late Campanian-early Maastrichtian sedimentary succession of the James Ross Basin (NG Sequence). Its major exposures occur at the James Ross and Vega islands. Several fossil-bearing localities have been identified in the SHIF providing a valuable fauna of invertebrates and vertebrates, and flora. Our study focuses on the vertebrate fauna recovered at Gamma and Cape Lamb members of the SHIF. The marine vertebrate assemblages include chondrichthyans, actinopterygians, and marine reptiles (elasmosaurid plesiosaurs and mosasaurs). A diverse terrestrial vertebrate assemblage has been reported being characterized by dinosaurs (sauropod, elasmarian ornithopods, nodosaurid ankylosaur, and a paravian theropod), pterosaurs and birds. Most SHIF dinosaurs share close affinities with penecontemporaneous taxa from southern South America, indicating that at least some continental vertebrates could disperse between southern South America and Antarctica during the Late Cretaceous. The Snow Hill Island Formation provides the most diverse Late Cretaceous marine and continental faunas from Antarctica. The present study summarizes previous and new vertebrate findings with the best actualized stratigraphical framework, providing a more complete fauna association and analyzing further perspectives.
- 1 January 2022
- Published by FapUNIFESP (SciELO) in Anais da Academia Brasileira de Ciências
- Vol. 94 (suppl 1) , e20211142
Open AccessLate Campanian-Early Maastrichtian Vertebrates From The James Ross Basin, West Antarctica: Updated Synthesis, Biostratigraphy, And Paleobiogeography
The Snow Hill Island Formation (SHIF; late Campanian - early Maastrichtian) crops out in the northeast of the Antarctic Peninsula and constitutes the basal part of the late Campanian-early Maastrichtian sedimentary succession of the James Ross Basin (NG Sequence). Its major exposures occur at the James Ross and Vega islands. Several fossil-bearing localities have been identified in the SHIF providing a valuable fauna of invertebrates and vertebrates, and flora. Our study focuses on the vertebrate fauna recovered at Gamma and Cape Lamb members of the SHIF. The marine vertebrate assemblages include chondrichthyans, actinopterygians, and marine reptiles (elasmosaurid plesiosaurs and mosasaurs). A diverse terrestrial vertebrate assemblage has been reported being characterized by dinosaurs (sauropod, elasmarian ornithopods, nodosaurid ankylosaur, and a paravian theropod), pterosaurs and birds. Most SHIF dinosaurs share close affinities with penecontemporaneous taxa from southern South America, indicating that at least some continental vertebrates could disperse between southern South America and Antarctica during the Late Cretaceous. The Snow Hill Island Formation provides the most diverse Late Cretaceous marine and continental faunas from Antarctica. The present study summarizes previous and new vertebrate findings with the best actualized stratigraphical framework, providing a more complete fauna association and analyzing further perspectives.
- Open Access
- 25 November 2020
- Published by Springer Nature in Communications Biology
- Vol. 3 (1) , 1-9
Persistent biotic interactions of a Gondwanan conifer from Cretaceous Patagonia to modern Malesia
Many plant genera in the tropical West Pacific are survivors from the paleo-rainforests of Gondwana. For example, the oldest fossils of the Malesian and Australasian coniferAgathis(Araucariaceae) come from the early Paleocene and possibly latest Cretaceous of Patagonia, Argentina (West Gondwana). However, it is unknown whether dependent ecological guilds or lineages of associated insects and fungi persisted on Gondwanan host plants likeAgathisthrough time and space. We report insect-feeding and fungal damage on PatagonianAgathisfossils from four latest Cretaceous to middle Eocene floras spanning ca. 18 Myr and compare it with damage on extantAgathis. Very similar damage was found on fossil and modernAgathis, including blotch mines representing the first known Cretaceous–Paleogene boundary crossing leaf-mine association, external foliage feeding, galls, possible armored scale insect (Diaspididae) covers, and a rust fungus (Pucciniales). The similar suite of damage, unique to fossil and extantAgathis, suggests persistence of ecological guilds and possibly the component communities associated withAgathissince the late Mesozoic, implying host tracking of the genus across major plate movements that led to survival at great distances. The living associations, mostly made by still-unknown culprits, point to previously unrecognized biodiversity and evolutionary history in threatened rainforest ecosystems.
- 25 November 2020
- Published by Springer Nature in Communications Biology
- Vol. 3 (1) , 1-9
Open AccessPersistent biotic interactions of a Gondwanan conifer from Cretaceous Patagonia to modern Malesia
Many plant genera in the tropical West Pacific are survivors from the paleo-rainforests of Gondwana. For example, the oldest fossils of the Malesian and Australasian coniferAgathis(Araucariaceae) come from the early Paleocene and possibly latest Cretaceous of Patagonia, Argentina (West Gondwana). However, it is unknown whether dependent ecological guilds or lineages of associated insects and fungi persisted on Gondwanan host plants likeAgathisthrough time and space. We report insect-feeding and fungal damage on PatagonianAgathisfossils from four latest Cretaceous to middle Eocene floras spanning ca. 18 Myr and compare it with damage on extantAgathis. Very similar damage was found on fossil and modernAgathis, including blotch mines representing the first known Cretaceous–Paleogene boundary crossing leaf-mine association, external foliage feeding, galls, possible armored scale insect (Diaspididae) covers, and a rust fungus (Pucciniales). The similar suite of damage, unique to fossil and extantAgathis, suggests persistence of ecological guilds and possibly the component communities associated withAgathissince the late Mesozoic, implying host tracking of the genus across major plate movements that led to survival at great distances. The living associations, mostly made by still-unknown culprits, point to previously unrecognized biodiversity and evolutionary history in threatened rainforest ecosystems.
- Open Access
- 8 October 2020
- Published by Cambridge University Press (CUP) in Paleobiology
- Vol. 46 (4) , 445-469
Cretaceous–Paleogene plant extinction and recovery in Patagonia
The Cretaceous–Paleogene (K/Pg) extinction appears to have been geographically heterogeneous for some organismal groups. Southern Hemisphere K/Pg palynological records have shown lower extinction and faster recovery than in the Northern Hemisphere, but no comparable, well-constrained Southern Hemisphere macrofloras spanning this interval had been available. Here, macrofloral turnover patterns are addressed for the first time in the Southern Hemisphere, using more than 3500 dicot leaves from the latest Cretaceous (Maastrichtian) and the earliest Paleocene (Danian) of Argentine Patagonia. A maximum ca. 90% macrofloral extinction and ca. 45% drop in rarefied species richness is estimated across the K/Pg, consistent with substantial species-level extinction and previously observed extirpation of host-specialized leaf mines. However, prior palynological and taxonomic studies indicate low turnover of higher taxa and persistence of general floral composition in the same sections. High species extinction, decreased species richness, and homogeneous Danian macrofloras across time and facies resemble patterns often observed in North America, but there are several notable differences. When compared with boundary-spanning macrofloras at similar absolute paleolatitudes (ca. 50°S or 50°N) from the Williston Basin (WB) in the Dakotas, both Maastrichtian and Danian Patagonian species richnesses are higher, extending a history of elevated South American diversity into the Maastrichtian. Despite high species turnover, our analyses also reveal continuity and expansion of leaf morphospace, including an increase in lobed and toothed species unlike the Danian WB. Thus, both Patagonian and WB K/Pg macrofloras support a significant extinction event, but they may also reflect geographically heterogeneous diversity, extinction, and recovery patterns warranting future study.
- 8 October 2020
- Published by Cambridge University Press (CUP) in Paleobiology
- Vol. 46 (4) , 445-469
Open AccessCretaceous–Paleogene plant extinction and recovery in Patagonia
The Cretaceous–Paleogene (K/Pg) extinction appears to have been geographically heterogeneous for some organismal groups. Southern Hemisphere K/Pg palynological records have shown lower extinction and faster recovery than in the Northern Hemisphere, but no comparable, well-constrained Southern Hemisphere macrofloras spanning this interval had been available. Here, macrofloral turnover patterns are addressed for the first time in the Southern Hemisphere, using more than 3500 dicot leaves from the latest Cretaceous (Maastrichtian) and the earliest Paleocene (Danian) of Argentine Patagonia. A maximum ca. 90% macrofloral extinction and ca. 45% drop in rarefied species richness is estimated across the K/Pg, consistent with substantial species-level extinction and previously observed extirpation of host-specialized leaf mines. However, prior palynological and taxonomic studies indicate low turnover of higher taxa and persistence of general floral composition in the same sections. High species extinction, decreased species richness, and homogeneous Danian macrofloras across time and facies resemble patterns often observed in North America, but there are several notable differences. When compared with boundary-spanning macrofloras at similar absolute paleolatitudes (ca. 50°S or 50°N) from the Williston Basin (WB) in the Dakotas, both Maastrichtian and Danian Patagonian species richnesses are higher, extending a history of elevated South American diversity into the Maastrichtian. Despite high species turnover, our analyses also reveal continuity and expansion of leaf morphospace, including an increase in lobed and toothed species unlike the Danian WB. Thus, both Patagonian and WB K/Pg macrofloras support a significant extinction event, but they may also reflect geographically heterogeneous diversity, extinction, and recovery patterns warranting future study.
- 1 May 2020
- Published by Elsevier in Cretaceous Research
- Vol. 109
Palynology from the Cenomanian Mata Amarilla Formation, southern Patagonia, Argentina
- 1 May 2020
- Published by Elsevier in Cretaceous Research
- Vol. 109
Palynology from the Cenomanian Mata Amarilla Formation, southern Patagonia, Argentina