Contributions towards whole-plant reconstructions of Dicroidium plants (Umkomasiaceae) from the Permian of Jordan

Blomenkemper, Patrick; Kerp, Hans; Hamad, Abdalla Abu; Bomfleur, Benjamin

doi:10.1016/j.revpalbo.2020.104210

Cited by 15 publications

(13 citation statements)

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“…Adaptation to these disturbance-prone environments may have played a key role in the ability of the plants to cope with the dramatic changes in the course of the Permian-Triassic transition and their success during the Triassic and Jurassic. Unlike highly competitive groups preferring stable humid conditions, such as gigantopterids and glossopterids that vanish from the fossil record at or shortly after the P/T boundary, Bennettitales thrived throughout the Triassic reaching a global distribution during the Late Triassic, similar to other groups whose earliest bona fide records, i.e., the Corystospermales and podocarpacean conifers, have been documented from these environments (Abu Hamad et al, 2008;Blomenkemper et al, 2018Blomenkemper et al, , 2020. The Bennettitales and other plants growing in periodically drier environments show a remarkable resilience toward the dramatic turnovers (Bomfleur et al, 2018) that affected the terrestrial and marine fauna so strongly (e.g., Erwin, 1999;Sahney and Benton, 2008).…”

Section: Discussionmentioning

confidence: 95%

“…Sediments of the Umm Irna Formation were deposited in a fluviolacustrine lowland setting under strongly seasonal conditions (e.g., Bandel and Khoury, 1981;Stephenson and Powell, 2013). The formation has yielded a peculiar admixture of plant taxa, including typical late Paleozoic and Mesozoic taxa as well as typical representatives from different floral provinces of Pangea (Mustafa, 2003;Kerp et al, 2006;Abu Hamad et al, 2008Blomenkemper et al, 2018Blomenkemper et al, , 2019Blomenkemper et al, , 2020Zavialova et al, 2021). Bennettitalean fossils have been recovered from three fossil localities in particular:…”

Section: Methodsmentioning

confidence: 99%

“…Moreover, some taxa appear to have persisted much longer than originally thought, such as the walchian conifer Otovicia hypnoides, a very long-ranging species that has its last appearance in the upper Permian of Jordan. The first occurrences of at least three major lineages of gymnospermous plants that were traditionally regarded as typically Mesozoic were reported from the upper Permian of Jordan (Kerp et al, 2006;Abu Hamad et al, 2008Blomenkemper et al, 2018Blomenkemper et al, , 2020. Most common are corystosperms of which first foliage assignable to Dicroidium was described (Kerp et al, 2006;Abu Hamad et al, 2008, and recently also fertile organs (Blomenkemper et al, 2020).…”

Section: Introductionmentioning

confidence: 99%

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Bennettitalean Leaves From the Permian of Equatorial Pangea—The Early Radiation of an Iconic Mesozoic Gymnosperm Group

et al. 2021

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Bennettitaleans are an extinct group of gymnosperms that are among the most iconic plants of Earth’s vegetation during the Mesozoic Era. The sudden appearance and rise to dominance of the Bennettitales during the Triassic remains a mystery. Leaf fossils similar to typical bennettitalean foliage occur in late Paleozoic deposits worldwide, but bennettitalean foliage can be identified with certainty only in case the fossils are sufficiently well-preserved to show epidermal features. So far, the characteristic stomatal architecture of the group has never been systematically documented in these putative Paleozoic remains. Here, we present well-preserved bennettitalean leaves from Permian deposits in two widely separated regions of equatorial Pangea. Two species of cuticle-bearing leaf compressions from the late Permian Umm Irna Formation, Jordan, are here formally described as Pterophyllum pottii Bomfleur et Kerp sp. nov. and Nilssoniopteris jogiana Blomenkemper et Abu Hamad sp. nov. Moreover, bulk maceration of samples from the Umm Irna Formation yielded six additional types of dispersed bennettitalean cuticles that are here informally described. In addition, the Cisuralian (early Permian) uppermost part of the Upper Shihhotse Formation exposed at the Palougou section in Shanxi Province, China, has yielded the oldest unambiguous bennettitalean fossils known to date; they consist of fragments of entire-margined leaves with well-preserved cuticles that we assign to Nilssoniopteris shanxiensis Bäumer, Backer et Wang sp. nov. Unlike the characteristic puzzle-patterned cuticles typical of many Jurassic and Cretaceous bennettites, the cuticles of these Permian bennettitalean remains show non-sinuous anticlinal walls, greater variety in stomatal orientation, and rare occurrence of transversely divided subsidiary cells—features that have until now almost exclusively been documented from the hitherto oldest cuticle-bearing Triassic bennettitalean material. Finally, the taxonomic richness, disjunct distribution, and broad variety in macro- and micromorphological features in these Permian bennettitalean remains lead us to suspect that the origin of the group will date back still further in time, and might in fact coincide with very early occurrences of Bennettitales-like foliage from the Pennsylvanian and Cisuralian, such as Pterophyllum cottaeanum, P. eratum, or P. grandeuryi.

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Section: Discussionmentioning

confidence: 95%

Section: Methodsmentioning

confidence: 99%

Section: Introductionmentioning

confidence: 99%

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Bennettitalean Leaves From the Permian of Equatorial Pangea—The Early Radiation of an Iconic Mesozoic Gymnosperm Group

et al. 2021

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“…Mesozoic gymnospermous plants bearing seeds within a "cupule" are commonly referred to as "pteridosperms" or "Mesozoic seed ferns", terms that reflect the traditional recognition that leaf and seed-bearing structures of these plants are at least superficially similar to those of Paleozoic pteridosperms (Taylor et al, 2006(Taylor et al, , 2009a. While there is some emerging evidence for phylogenetic continuity for some of these plants across the Permian-Triassic boundary (Kerp et al, 2001;Hamad et al, 2017;Blomenkemper et al, 2018Blomenkemper et al, , 2020Anderson et al, 2019), most groups are unlikely to share close affinities with Paleozoic seed plants (Figure 21). Instead, as is evident from lagerstätte like Apple Bay, many lineages of Mesozoic gymnosperms "experimented" with seedenclosing structures.…”

Section: Morphology Anatomy and Homologies Among Seed-bearing Structu...mentioning

confidence: 99%

A novel cupulate seed plant, Xadzigacalix quatsinoensis gen. et sp. nov., provides new insight into the Mesozoic radiation of gymnosperms

Klymiuk

Rothwell

Stockey

2022

American J of Botany

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Premise Anatomically preserved evidence for a novel clade of gymnosperms emphasizes diversity of seed plants immediately prior to the appearance of angiosperm fossils in the paleontological record. Methods Cupulate seeds from the Early Cretaceous Apple Bay locality (Vancouver Island) are described from serial cellulose acetate peels and three‐dimensional reconstruction. Phylogenetic context is assessed through the comparative analysis of gymnosperm seed producing fructifications and maximum parsimony analysis of a revised morphological data set for seed plant phylogeny. Results Xadzigacalix quatsinoensis gen. et sp. nov. is characterized by an orthotropous ovule with an elongated micropyle and complex integument, enclosed within a radial cupule. The micropylar canal is elongated; and the nucellus extends into the micropyle to seal the post pollination ovule. Except at the apex of the micropyle, the seed is completely enclosed by a parenchymatous cupule with ca. 20 axially elongated secretory ducts. The cupulate seed is produced upon a triangular woody stele, consisting of a parenchymatous pith surrounded by radially aligned tracheids. The stele produces three short terete traces that terminate within the base of the cupule as transfusion tissue at the seed chalaza. Conclusions Organography, vascularization, nature of the integument and nucellus, and configuration of the micropylar canal distinguish Xadzigacalix quatsinoensis from all other gymnosperm clades. Cladistic analyses suggest the new plant may have affinities with gnetophytes or angiosperms. These results are complemented with a critical re‐evaluation of ovulate structures for Mesozoic gymnosperms, providing new insight into plant diversity immediately antecedent to the explosive diversification of flowering plants.

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“…Nevertheless, the anatomical data obtained from this deposit are difficult to integrate systematically into the group's impression/compression fossil record. There have been only rare attempts to reconstruct umkomasialean plants as a whole (RETALLACK 1977;CRANE 1985;RETALLACK & DILCHER 1988;TAYLOR et al 2006;BLOMENKEMPER et al 2020).…”

Section: Table Of Contentsmentioning

confidence: 99%

Mummified Dicroidium (Umkomasiales) leaves and reproductive organs from the Upper Triassic of South Australia

Unverfärth¹,

McLoughlin²,

Bomfleur³

2022

palb

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The Leigh Creek Coal Measures incorporate unusually low-rank coals from the Upper Triassic of South Australia. Associated fluvio-lacustrine deposits contain well-preserved, partly mummified plant remains dominated by corystosperm seed ferns. The assemblage comprises seven species of Dicroidium, including D. odontopteroides, D. lineatum, D. dubium, D. zuberi, and Dicroidium spp. A, B and C, and associated reproductive organs, including various fragments of cupulate structures (Umkomasia sp. cf. U. quadripartita and Fanerotheca sp. cf. F. waldeckiformis) and pollen organs (Pteruchus africanus), all having excellent cuticle preservation. Based on a comprehensive analysis of more than 550 individual specimens, we (1) document diagnostic epidermal and cuticular features for foliage and reproductive organs, (2) provide an identification key for the Dicroidium species present, and (3) infer affiliations between reproductive organs and particular leaf species based on correspondence in epidermal anatomy and cuticle micromorphology and on mutual-occurrence data. Collectively, the Leigh Creek material contributes towards a more robust and realistic systematic classification of Umkomasiaceae, offers a rare chance for whole-plant reassembly of individual species, and refines reconstruction of the Dicroidiumdominated forest ecosystems in the middle to high latitudes of the Late Triassic greenhouse world.

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Contributions towards whole-plant reconstructions of Dicroidium plants (Umkomasiaceae) from the Permian of Jordan

Cited by 15 publications

References 51 publications

Bennettitalean Leaves From the Permian of Equatorial Pangea—The Early Radiation of an Iconic Mesozoic Gymnosperm Group

Bennettitalean Leaves From the Permian of Equatorial Pangea—The Early Radiation of an Iconic Mesozoic Gymnosperm Group

A novel cupulate seed plant, Xadzigacalix quatsinoensis gen. et sp. nov., provides new insight into the Mesozoic radiation of gymnosperms

Mummified Dicroidium (Umkomasiales) leaves and reproductive organs from the Upper Triassic of South Australia

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