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P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F-H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C-E, 1 cm; F, 100 μm; G and H, 200 μm; I-L, 5 mm; M, 2 mm; N, 10 μm.)
Source publication
Significance
There were two heterosporous lignophyte lineages of which only one, the seed plants, survived the Permian–Triassic mass extinction. Based on exceptionally complete fossil trees from a 300-My-old volcanic ash, the enigmatic Noeggerathiales are now recognized as belonging to the other lineage. They diversified alongside the primary seed...
Contexts in source publication
Context 1
... fossils were collected from a single 66-cm-thick volcanic ash bed in the Chinese "vegetational Pompeii" from the Taiyuan Formation at Wuda open coalmine, Inner Mongolia (14, 15) (SI Appendix, Geological Information, Materials). The ash preserved in situ the morphology and anatomy of plants in exquisite detail ( Fig. 1 A-N) and has been dated to 298.34 ± 0.09 Ma during the Asselian stage of the early Permian (16). ...
Context 2
... Appendix, Figs. S3-S8) and megaphyllous leaves (SI Appendix, Figs. S9-S14) are common in the assemblage. The species P. wuhaia is based on >200 specimens, including >10 specimens that were preserved with intact crowns containing both leaves and fertile shoots in organic attachment to the stems, demonstrating that they belong to a single species (Fig. 1A and SI Appendix, Figs ...
Context 3
... (Fig. 1A and SI Appendix, Repository-All specimens are deposited at Nanjing Institute of Geology and Paleontology, Chinese Academy of ...
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... The holotype is 160 cm long and 60 cm wide ( Fig. 1A and SI Appendix, Fig. S2) and comprises a stem with apical crown of helically attached megaphyllous leaves and pseudostrobili. The crown has three parts: a 13-cm-long basal region bearing leaves, a middle 23-cm-long fertile zone, and apically a 6-cm-long region bearing more leaves. Below the crown, the stem is naked, extending for >32 ...
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... pseudostrobili show four to five rows of sporangia with rows arranged parallel to the rachis ( Fig. 1 I and K and SI Appendix, Figs. S5-S8). As the specimens have been affected by postmortem compression (SI Appendix, Figs. S5-S8), we consider that there must have been a row of sporangia on either side that was laterally flattened; we estimate that a complete pseudostrobilus comprised 8 or 10 sporangia per whorl. Sporophylls are ...
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... Figs. S5-S8), we consider that there must have been a row of sporangia on either side that was laterally flattened; we estimate that a complete pseudostrobilus comprised 8 or 10 sporangia per whorl. Sporophylls are proximally fused laterally to adjacent sporophylls, while distally they are free with a small heel and upturned apex (SI Appendix, Figs. S21D and S23E). There are 32 to 35 sporangia per row in the pseudostrobili depending on the length of the pseudostrobili, producing five sporangia per centimeter's length. Sporangia are sessile, attached to the sporophylls adaxial surface (Fig. 1 I-M). Megasporangia occur in a single vertical row on the pseudostrobilus axis ( Fig. 1 ...
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... to adjacent sporophylls, while distally they are free with a small heel and upturned apex (SI Appendix, Figs. S21D and S23E). There are 32 to 35 sporangia per row in the pseudostrobili depending on the length of the pseudostrobili, producing five sporangia per centimeter's length. Sporangia are sessile, attached to the sporophylls adaxial surface (Fig. 1 I-M). Megasporangia occur in a single vertical row on the pseudostrobilus axis ( Fig. 1 ...
Context 8
... are sessile, attached to the sporophylls adaxial surface (Fig. 1 I-M). Megasporangia occur in a single vertical row on the pseudostrobilus axis ( Fig. 1 K- ...
Context 9
... of the hormone auxin under gravity to trigger megasporangial expression (9). Microspores are of the Calamospora type; trilete circular, subcircular to oval, 30 to 40 μm in diameter, with straight rays of the trilete mark forming a labrum extending 3/4 or the whole radius a laevigate exine that forms irregular folds usually parallel to the margin (Fig. 1N and SI Appendix, Fig. ...
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... Description. P. wuhaia J. Wang et al. is reconstructed as a small, slender, unbranched palm-like tree in Fig. 2, two to many meters high. Its major characteristics are illustrated by a diagrammatic reconstruction as shown in Fig. 3. The details of its stem anatomy are consistent with that of progymnosperms and early seed plants (SI Appendix, Figs. ...
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Citations
... Leary, 1980;Wang et al. 2004;Taylor et al. 2009). Recently, the finding and description of whole specimens found within a Permian ash-tuff with articulated foliage, (pseudo-like)strobili and stem with anatomical preservation, allowed the elucidation of the Noeggerathiales as members of the progymnosperms clade, a sister group to the seed plants (Wang et al. 2021;Yang et al. 2023). ...
Noeggerathiales were until recently a group of plants with uncertain systematic position that existed in the Carboniferous and Permian times. Recent discoveries classify them as heterosporous progymnosperms. Despite the discovery of additional specimens, the group still remains highly artificial because their reproductive organs are rarely preserved in organic connection. Within the Carboniferous of Iberian Massif, the noeggerathialeans are poorly represented. Here, we describe Palaeopteridium andrenelii sp. nov. from the uppermost Carboniferous of Portugal. This is the second representative of Noeggerathiales reported in the Portuguese Carboniferous after Carlos Teixeira have described the noeggerathialean Rhacopteris gomesiana in the 1940s from Douro Carboniferous Basin (Stephanian C/lower Gzhelian, Upper Pennsylvanian). Palaeopteridium andrenelii was found in upper Asturian (upper Moscovian, Middle Pennsylvanian) strata from the classical Westphalian outcrops of Ervedosa, located in the region of Alto da Serra (Fânzeres), Gondomar, in northwestern Portugal. Two reproductive structures are associated with the frond of the new fossil species. Although not organically linked, both structures could belong to parent plant (frond) and represent possible detached macrosporangia. This reenforces the Palaeopteridium as a noeggerathialean and the first reproductive structures found for this genus.
... Progymnosperm fossils at CUVA are currently limited to wood of the form genus Callixylon reported from the Bedford Formation and a possible noeggerathialean leaf specimen observed in the Cuyahoga Group ( Figure 12). Plants of the order Noeggerathiales have been historically enigmatic; however, recent well-preserved fossils support the placement of this clade as derived progymnosperms (Wang et al. 2021). Given the rarity of noeggerathialean fossils in the Carboniferous Euramerican floral realm (Pfefferkorn and Wang 2016), the specimen observed in the Mississippian sandstone of CUVA would hold significance, if the identification is accurate. ...
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... Progymnosperms are a distinct group with a fern-like morphology and gymnosperm inner structure (Beck 1960) and a geological history ranging from the Middle Devonian as far as the Permian (Wang et al. 2021). Progymnosperms are likely to inhabit highland areas or locations farther away from the coast due to their higher water-use efficiency (Wan et al. 2019), and their dispersal is highly dependent on wind. ...
Dispersal, whether active or passive, plays a crucial role in biogeography by facilitating the movement of propagules away from their original location. Botanical geographical zonation, resulting from the co-evolution of plants and their environment, has been established since the remarkable plant diversification during the Devonian Period (c. 419–359 Ma). However, a significant knowledge gap exists in understanding plant dispersal between living and fossil organisms due to the rarity of opportunities for tracing plant dispersal in geological history. In this study, we present evidence of two plant dispersal routes and verify their occurrence through the examination of geographical zonation, changes in plant diversity, and latitudinal and longitudinal gradients during the Devonian. We analyse global occurrence data from widely-distributed and extensively-studied Devonian plants. The two dispersal routes, namely clockwise and anticlockwise, connect the South China and Euramerica–Siberia realms. These routes clearly demonstrate inland and inter-land dispersal models, closely linked to Devonian sea–land topography and dispersal vectors such as wind and ocean currents. Moreover, these models probably apply to all Devonian plants. Our comprehensive synthesis of plant dispersal in deep time reveals that propagule diversity and dispersal vectors have progressively increased and become more complex over time, facilitating plant colonization and diversity changes. Importantly, our study unveils the dispersal models of fossil plants, demonstrating the equivalent models observed in extant plants that have been established since the Devonian Period.
... These differences in cone architecture are useful in inferring relationships between different plant groups. For example, strobili with affinity to Calamitales and Sphenophyllales (extinct horsetail relatives) exhibit whorled sporophylls (Hoskins and Cross, 1943;Baxter, 1950;Stein et al., 1984), whereas those with affinity to Noeggerathiales (the freesporing sister clade to the seed plants; Wang et al., 2021) exhibit alternately arranged sporophylls (Leary, 1973;Leary and Pfefferkorn, 1977;Gao and Zodrow, 1990). Others, such as the rare Tetraphyllostrobus Gao et Zodrow, which has been described as exhibiting decussate sporophyll arrangement (Gao and Zodrow, 1990), are enigmatic and do not conform to the architecture of any Paleozoic plant group (Friis, 1992). ...
Premise
We studied the 3D morphology of a small, well‐preserved cone from the Pennsylvanian Mazon Creek Lagerstätte to characterize its structure and determine its systematic affinity. Previously tentatively assigned to the enigmatic Tetraphyllostrobus , we show that it differs in key respects from that genus as described.
Methods
We systematically compared the new fossil with relevant Paleozoic cone genera and employed advanced imaging techniques, including scanning electron microscopy, Airyscan confocal super‐resolution microscopy, optical microscopy, and X‐ray microcomputed tomography to visualize and reconstruct the fossil cone in 3D.
Results
The analyses demonstrate unequivocally that the sporophylls of the new Mazon Creek cone are arranged in whorls of six and have characters typical of Sphenophyllales, including epidermal cells with undulatory margins and in situ spores assignable to Columinisporites . The combination of characters, including sporophyll arrangement, anatomy, and spore type, supports the establishment of Hexaphyllostrobus kostorhysii gen. et sp. nov. within Sphenophyllales. Furthermore, we show that Tetraphyllostrobus , although originally described as possessing smooth monolete spores, actually possesses Columinisporites ‐type spores, indicating that it, too, was most likely a sphenophyll.
Conclusions
The recognition of Hexaphyllostrobus contributes to our knowledge of Pennsylvanian sphenophyll diversity, and in particular increases the number of species with in situ Columinisporites ‐type spores. Attribution of Hexaphyllostrobus to Sphenophyllales calls into question current interpretations of Tetraphyllostrobus suggesting that future research on better‐preserved macrofossil material may demonstrate a sphenophyllalean relationship.
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... The order Noeggerathiales is an enigmatic and extinct plant group in the Late Palaeozoic on Cathaysia and Euramerica continents. Since it was first proposed in 1931, around 50 species of 20 genera have been enrolled [5][6][7]. However, the systematic affinity of this group was poorly understood until the stem anatomy of Paratingia Zhang was studied [7]. ...
... Since it was first proposed in 1931, around 50 species of 20 genera have been enrolled [5][6][7]. However, the systematic affinity of this group was poorly understood until the stem anatomy of Paratingia Zhang was studied [7]. Up to now, growth habit has been reported for only four species, namely Noeggerathia foliosa [8], Paratingia wudensis [9], Paratingia wuhaia [7], and Tingia unita [10,11], of which only Paratingia wuhaia is known for stem anatomy [7], and accordingly it has been concluded ...
... However, the systematic affinity of this group was poorly understood until the stem anatomy of Paratingia Zhang was studied [7]. Up to now, growth habit has been reported for only four species, namely Noeggerathia foliosa [8], Paratingia wudensis [9], Paratingia wuhaia [7], and Tingia unita [10,11], of which only Paratingia wuhaia is known for stem anatomy [7], and accordingly it has been concluded ...
Simple Summary
The systematic position of Noeggerathiales was long uncertain until the whole plant species Paratingia wuhaia was restored and proved to belong to progymnosperms due to its spore-producing fertile organ and secondary wood producing a large stem. However, whether Tingia Halle as the most diversified genus in Noeggerathiales belongs to progymnosperms has yet remained uncertain as the anatomy of the main stem of this plant is unknown, that is, whether the anatomy of the main stem with the anatomical characteristics of gymnosperms woods remains uncertain. Here, the stem anatomy of Tingia unita is presented based on fossil materials from the early Permian Wuda Tuff Flora in Wuda Coalfield, Wuhai City, Inner Mongolia, China. The well-developed secondary wood, plus previously accumulated evidence of the spore-bearing nature of this plant, confirms that this genus belongs to progymnosperms. As such, the genera Tingia and Paratingia are all certainly progymnosperms in affinity.
Abstract
Tingia Halle, a representative genus of the Cathaysia Flora, has been studied for nearly 100 years, being a small heterosporous tree based on the gross morphology of Tingia unita. However, the systematic affinity of Tingia is uncertain. Now, a number of well-preserved fossils of T. unita from the Taiyuan Formation of Lower Permian in Wuda Coalfield, Wuhai City, Inner Mongolia facilitates an examination of wood anatomy. The stem anatomy of T. unita shows parenchymatous pith, endarch primary xylem, pycnoxylic secondary xylem, and cortex, typically a type of gymnosperm wood, which taken together with pteridophytic reproduction, certainly evidences that Tingia Halle is a progymnosperm. In addition, Tingia together with Paratingia provide strong evidence to link the Noeggerathiales with progymnosperms.
... They also reveal partial permineralization of the internal anatomy. Collectively, Wuda plant fossils facilitate in-depth systematic studies (e.g., Wang J. et al., 2021a). Based on Wuda specimens, Zhang et al. (2008) made a conceptual reconstruction of the tripinnate Pecopteris lativenosa pinnae by statistical analysis, revealing the high intraspecific morphological variation of the frond. ...
Pecopteris (Brongniart) Sternberg is a morphological fossil genus with a cosmopolitan distribution during the Permo–Carboniferous, of which most species belong to the marattialean and or “filicalean” ferns or less frequently to pteridosperms. The taxonomic affinity is uncertain unless distinctive reproductive organs or rachis anatomy is known. Among the numerous species of Pecopteris in the Permian Wuda Tuff Flora, Inner Mongolia, China, Pecopteris lativenosa Halle has been considered as a pteridosperm based on its larger variation of frond morphology, although evidence of fertile organs or rachises are unknown. Newly discovered specimens of this species from the Wuda Tuff Flora are mostly impression fossils. However, some have partially permineralized petioles and have fronds in organic connection to Caulopteris-type stem. Fronds are tripinnate; penultimate pinnae lengths are variable as are ultimate pinnae lengths. Pinnules are oval and thick, and their midveins are straight and are of stable thickness. Lateral veins are dense, bifurcating 1–3 times. Petiole and rachises are anatomically preserved and show sclerenchyma, fundamental tissue and vascular tissue. Vascular bundles of petioles and rachises are C-shaped with two laterally inrolled ends, which are typically stewartiopterid petiole/rachis of Marattiales. These features allow us to assign Pecopteris lativenosa to the late Paleozoic marattialean family Psaroniaceae.
... Lignophytes are characterized by their bifacial vascular cambium, and encompass both progymnosperms and spermatophytes (Crane, 1985;Steemans et al., 2012;Momont et al., 2016;Decombeix et al., 2019). Progymnosperms (Aneurophytales, Archaeopteridales, Protopityales, and probably Noeggerathiales) are generally considered to be a sister group to seed plants (Crane et al., 2004;Hilton and Bateman, 2006;Niklas and Crepet, 2020;Wang et al., 2021;Figs. 1, 4). ...
... 1, 4). The Aneurophytales are homosporous and are assumed to be a sister clade to the heterosporous Archaeopteridales and spermatophytes, although these relationships are not yet resolved Toledo et al., 2018;Wang et al., 2021). Spermatophytes are strongly resolved as a monophyletic group Crane, 1997a, 1997b;Magallón and Hilu, 2009;Toledo et al., 2018;Fig. ...
During the mid-Palaeozoic, vascular land plants (i.e., tracheophytes) underwent a great radiation that triggered the development of the land biosphere – the so-called Silurian–Devonian terrestrial revolution. However, little is known about how different plant groups impacted this process. A newly constructed dataset of plant macrofossil genera is used to characterize the tempo and mode of development of Silurian–Devonian vegetation and how it spread out over subaerial habitats. Important fluctuations of diversity and evolutionary rates of vegetation are linked to the diversity dynamics of particular tracheophyte groups. Despite a general increase of taxonomic richness through the Devonian, there was a clear stepwise pattern of origination and extinction events that resulted in the main floral transitions over time, such as the change to a forested landscape. To test if sampling bias may be affecting the observed diversity patterns, the latter were compared with the number of plant macrofossil localities as a proxy for sampling effort. This suggested a highly significant correlation between observed diversity and sampling effort, but it was not homogeneous, suggesting that at least some diversity fluctuations have a potential biological explanation. The sampling-corrected pattern of standing diversity suggests a clear increase of plant richness in the Pragian (Early Devonian) and Givetian (Middle Devonian), which may be related to the early expansion of the tracheophyte clades and the initial diversification of forested ecosystems, respectively. Further works should be focused on elucidate the impact of rock record on our understanding of Devonian plant diversification.
... The Permo-Triassic mass extinction, the greatest biotic crisis of the Phanerozoic, involved the loss of many groups of plants and animals (Hallam and Wignall, 1997;Bambach et al., 2004;Stanley, 2016). Its impact on terrestrial ecosystems was severe, with peat-forming, lowland plant communities vanishing for tens of millions of years (Retallack et al., 1996) and many higher plant taxa, such as the glossopterid, gigantopterid and cordaitalean gymnosperms, and the noeggerathialeans, becoming extinct (Retallack, 1995;Zhang et al., 2016;Fielding et al., 2019;Mays et al., 2020;Wang et al., 2021). These losses broadly coincided with the eruption of the Siberian Traps. ...
In terrestrial settings, the Permo-Triassic mass extinction is commonly linked to major changes in sedimentological and climatic conditions that include a switch from meandering to braided fluvial systems and increased aridity. We examined the predominantly terrestrial strata of North China to reveal that, on the contrary, there was little substantial sedimentological change during the Permo-Triassic mass extinction. The crisis level occurs in the upper Sunjiagou Formation, where a range of environments, including low-sinuosity rivers, distally terminating alluvial fans, and mud-dominated coastal plains are recorded. A major sedimentological change occurred slightly higher, within the lowest Triassic, and involved a switch to braided fluvial and shallow lacustrine/fluviodeltaic conditions of the Liujiagou Formation. Later in the Early Triassic, fine-grained, playa lake, and alluvial plain facies became widespread (Heshanggou Formation). Paleosols are present throughout the basin fill and record a transition from Aridisols and Vertisols in the Late Permian to Inceptisols in the Early Triassic followed by gleyed Aridisols, Vertisols, and Inceptisols later in the Early Triassic. Previously reported aeolian facies from Lower Triassic strata were not encountered in this study and were unlikely to have occurred given the prevalence of lacustrine and fluvial conditions at this time. Overall, the North China Basin experienced an increase in seasonal rainfall in the latest Permian and a base-level rise that introduced marine conditions into the southwest of the basin. During the Early Triassic, seasonal or interannual rainfall was frequent, with strong evaporation in a hot climate. Overall, substantial sedimentological changes occurred in the terrestrial Permo-Triassic environments of North China, but despite earlier claims from studies in North China and elsewhere, there was no abrupt transition in fluvial styles around the Permo-Triassic mass extinction.
... The Wuda Tuff Flora (also known as "vegetational Pompeii") is preserved between local coal seams No. 6 and No. 7 in a volcanic tuff bed that has undergone post-depositional alteration . The fossil plants preserved therein represent a swamp forest buried in situ by a single volcanic eruption (Wang et al., 2012), as evidenced by numerous erect stumps and fallen trees (e.g., Wan and Wang, 2020;Wang et al., 2009aWang et al., , 2009bWang et al., , 2021aWang et al., , 2021bWang et al., 2021;Liu et al., 2021;Zhou et al., 2021b). High-precision U-Pb dating of zircons established the absolute age of the tuff as 298.34 ± 0.09 Ma (Schmitz et al., 2021) in the earliest Permian. ...
Botryopteridaceae, characterized by an omega-shaped foliar trace, is one of the best known late Paleozoic fern families with frequent occurrence in the Carboniferous of Euramerica and recent recognition in Permian deposits of Gondwana and Cathaysia. Diodonopteris is a recently established botryopterid genus based on coal-ball material from the lower Permian Taiyuan Formation of Shanxi Province, China. Here, a new species of Diodonopteris is described based on an anatomically preserved specimen from the well-known early Permian Wuda Tuff Flora in the Wuda Coalfield of Inner Mongolia, China. Diodonopteris virgulata sp. nov. is similar to the type species D. gracilis in the marginal position of protoxylem groups but differs from the latter in having a simpler cortex and a more expanded xylem strand. The two known Diodonopteris species share several characteristics considered ancestral among botryopterids, including strand shape, mode of pinna trace divergence, weak pinnule lamination, and a laminar wing on the rachis, as is consistent with Diodonopteris sitting outside any of the derived sublineages within Botryopteris. D. virgulata is confirmed to be a climber by direct preservation with the host. Moreover, the host plant is also evidenced to be a climber. The repeated preservation of the dual-climbing phenomenon in this swamp forest signals the complexity of early Permian ecosystems.
