Guazia, the earliest ovule without cupule but with unique integumentary lobes.

The earliest ovules in the Late Devonian (Famennian) are surrounded by a cupule that is involved in both protection and pollination, and generally have free integumentary lobes. Here we report a new taxon from the Famennian of China, Guazia dongzhiensis gen. et sp. nov. The terminally borne ovule is apparently acupulate (without cupule) and has four radially arranged wing-like integumentary lobes that are extensively fused, and folded lengthwise and inwards. Guazia provides evidence that not all Devonian seeds possess a cupule and it increases their diversity in integumentary lobes. This genus also suggests that the integuments develop new functions, probably including wind dispersal at the expense of the cupules.

The Most Extensive Devonian Fossil Forest with Small Lycopsid Trees Bearing the Earliest Stigmarian Roots.

Since the Late Paleozoic, forests have become distributed worldwide and significantly changed the Earth's climate and landscapes, but the record of forests is rare in the Devonian (419-359 Ma in age) when they first appeared. From the Upper Devonian (Famennian with the age of 372-359 Ma) of Xinhang, Anhui, China, we report a very large in situ forest, which includes locally dense stands of lycopsid plants. The Xinhang forest is monospecific with a small tree lycopsid Guangdedendron gen. nov., probably dioecious with monocarpic reproduction. The plant shows the earliest stigmarian rooting system typical of giant tree lycopsids dominating Carboniferous forests. It colonizes coastal clastic wetlands that were influenced by floods. This significantly increases the paleogeographical coverage of in situ Devonian forests, and contributes to our understanding of atmospheric CO2 decline and coastal consolidation.

Further study of Late Devonian seed plant Cosmosperma polyloba: its reconstruction and evolutionary significance.

BACKGROUND: The earliest seed plants in the Late Devonian (Famennian) are abundant and well known. However, most of them lack information regarding the frond system and reconstruction. Cosmosperma polyloba represents the first Devonian ovule in China and East Asia, and its cupules, isolated synangiate pollen organs and pinnules have been studied in the preceding years. RESULTS: New fossils of Cosmosperma were obtained from the type locality, i.e. the Leigutai Member of the Wutong Formation in Fanwan Village, Changxing County, Zhejiang Province, South China. The collection illustrates stems and fronds extensively covered in prickles, as well as fertile portions including uniovulate cupules and anisotomous branches bearing synangiate pollen organs. The stems are unbranched and bear fronds helically. Fronds are dimorphic, displaying bifurcate and trifurcate types, with the latter possibly connected to fertile rachises terminated by pollen organs. Tertiary and quaternary rachises possessing pinnules are arranged alternately (pinnately). The cupule is uniovulate and the ovule has four linear integumentary lobes fused in basal 1/3. The striations on the stems and rachises may indicate a Sparganum-type cortex. CONCLUSIONS: Cosmosperma further demonstrates diversification of frond branching patterns in the earliest seed plants. The less-fused cupule and integument of this plant are considered primitive among Devonian spermatophytes with uniovulate cupules. We tentatively reconstructed Cosmosperma with an upright, semi-self-supporting habit, and the prickles along stems and frond rachises were interpreted as characteristics facilitating supporting rather than defensive structures.

Earliest filter-feeding pterosaur from the Jurassic of China and ecological evolution of Pterodactyloidea.

Pterosaurs were a unique clade of flying reptiles that were contemporaries of dinosaurs in Mesozoic ecosystems. The Pterodactyloidea as the most species-diverse group of pterosaurs dominated the sky during Cretaceous time, but earlier phases of their evolution remain poorly known. Here, we describe a 160 Ma filter-feeding pterosaur from western Liaoning, China, representing the geologically oldest record of the Ctenochasmatidae, a group of exclusive filter feeders characterized by an elongated snout and numerous fine teeth. The new pterosaur took the lead of a major ecological transition in pterosaur evolution from fish-catching to filter-feeding adaptation, prior to the Tithonian (145-152 Ma) diversification of the Ctenochasmatidae. Our research shows that the rise of ctenochasmatid pterosaurs was followed by the burst of eco-morphological divergence of other pterodactyloid clades, which involved a wide range of feeding adaptations that considerably altered the terrestrial ecosystems of the Cretaceous world.

Belowground rhizomes in paleosols: The hidden half of an Early Devonian vascular plant.

The colonization of terrestrial environments by rooted vascular plants had far-reaching impacts on the Earth system. However, the belowground structures of early vascular plants are rarely documented, and thus the plant-soil interactions in early terrestrial ecosystems are poorly understood. Here we report the earliest rooted paleosols (fossil soils) in Asia from Early Devonian deposits of Yunnan, China. Plant traces are extensive within the soil and occur as complex network-like structures, which are interpreted as representing long-lived, belowground rhizomes of the basal lycopsid Drepanophycus The rhizomes produced large clones and helped the plant survive frequent sediment burial in well-drained soils within a seasonal wet-dry climate zone. Rhizome networks contributed to the accumulation and pedogenesis of floodplain sediments and increased the soil stabilizing effects of early plants. Predating the appearance of trees with deep roots in the Middle Devonian, plant rhizomes have long functioned in the belowground soil ecosystem. This study presents strong, direct evidence for plant-soil interactions at an early stage of vascular plant radiation. Soil stabilization by complex rhizome systems was apparently widespread, and contributed to landscape modification at an earlier time than had been appreciated.

Latisemenia longshania, gen. et sp. nov., a new Late Devonian seed plant from China.

The earliest known ovules in the Late Devonian (Famennian) are borne terminally on fertile branches and are typically enclosed in a cupule. Among these ovules are some that have terete integumentary lobes with little or no fusion. Here, we report a new taxon, Latisemenia longshania, from the Famennian of South China, which bears cupulate ovules that are terminal as well as opposite on the fertile axis. Each ovule has four broad integumentary lobes, which are extensively fused to each other and also to the nucellus. The cupule is uniovulate, and the five flattened cupule segments of each terminal ovule are elongate cuneate and shorter than the ovule. Associated but not attached pinnules are laminate and Sphenopteris-like, with an entire or lobate margin. Latisemenia is the earliest known plant with ovules borne on the side of the fertile axis and may foreshadow the diverse ovule arrangements found among younger seed plant lineages that emerge in the Carboniferous. Following the telome theory, Latisemenia demonstrates derived features in both ovules and cupules, and the shape and fusion of integumentary lobes suggest effective pollination and protection to the nucellus. Along with other recent discoveries from China, Latisemenia extends the palaeogeographic range of the earliest seed plants.

Leaf evolution in early-diverging ferns: insights from a new fern-like plant from the Late Devonian of China.

BACKGROUND AND AIMS: With the exception of angiosperms, the main euphyllophyte lineages (i.e. ferns sensu lato, progymnosperms and gymnosperms) had evolved laminate leaves by the Late Devonian. The evolution of laminate leaves, however, remains unclear for early-diverging ferns, largely represented by fern-like plants. This study presents a novel fern-like taxon with pinnules, which provides new insights into the early evolution of laminate leaves in early-diverging ferns. METHODS: Macrofossil specimens were collected from the Upper Devonian (Famennian) Wutong Formation of Anhui and Jiangsu Provinces, South China. A standard degagement technique was employed to uncover compressed plant portions within the rock matrix. KEY RESULTS: A new fern-like taxon, SHOUGANGIA BELLA GEN ET SP NOV: , is described and represents an early-diverging fern with highly derived features. It has a partially creeping stem with adventitious roots only on one side, upright primary and secondary branches arranged in helices, tertiary branches borne alternately or (sub)oppositely, laminate and usually lobed leaves with divergent veins, and complex fertile organs terminating tertiary branches and possessing multiple divisions and numerous terminal sporangia. CONCLUSIONS: Shougangia bella provides unequivocal fossil evidence for laminate leaves in early-diverging ferns. It suggests that fern-like plants, along with other euphyllophyte lineages, had independently evolved megaphylls by the Late Devonian, possibly in response to a significant decline in atmospheric CO2 concentration. Among fern-like plants, planate ultimate appendages are homologous with laminate pinnules, and in the evolution of megaphylls, fertile organs tend to become complex.

A Late Devonian fertile organ with seed plant affinities from China.

Seed plants underwent first major evolutionary radiation in the Late Devonian (Famennian), as evidenced by the numerous ovules described to date. However, the early pollen organs are underrepresented, so that their structure and evolution remain poorly known. Here we report a new taxon of pollen organ Placotheca minuta from the Late Devonian. The synangium consists of many basally and more or less laterally fused microsporangia borne on the margin of a pad. The prepollen is spherical and trilete. The appearance of Famennian synangia especially in Placotheca does not support the current understanding that the earliest pollen organs closely resembled the fructifications of the ancestral progymnosperms. Placotheca indicates earlier diversification of pollen organs than previously expected and is highly derived among the early pollen organs with trilete prepollen. It is suggested that, immediately after the origination of seed plants, pollen organs had evolved at a rapid rate, whereas their prepollen remained primitively spore-like.

Cosmosperma polyloba gen. et sp. nov., a seed plant from the Upper Devonian of South China.

Seed plants with ovules were abundant in the Late Devonian of Euramerica and they contribute significantly to our understanding of their early history. However, coeval ovules have been scarce in other regions of the world. Specimens of the seed plant Cosmosperma polyloba gen. et sp. nov. Wang et al. were recently obtained from the Upper Devonian (Famennian) Wutong Formation, at Fanwan Village, Changxing County, Zhejiang Province, China. This new seed plant has cupulate ovules, the uniovulate cupules with up to 16 distal segments and with minute spines on the outer surface, synangiate pollen organs bearing six to eight microsporangia fused only at the base, and planate and highly dissected pinnules in alternate arrangement. It differs from other Devonian seed plants mainly in the organization and position of the uniovulate and ornamented cupule, and in the highly dissected pinnules. Cosmosperma Wang et al. represents the first Devonian ovules recovered from China or eastern Asia and further illustrates the diversity of early spermatophytes. As for the Late Devonian seed plants, it is suggested that the pollen organs are synangiate and simple in organization, and the branches and leaves are generally planate.

Diversity dynamics of silurian-early carboniferous land plants in South china.

New megafossil and microfossil data indicate four episodes in the diversification of Silurian-Early Carboniferous land plants of South China, a relatively continuous regional record. Plant diversity increased throughout, but the rising curve was punctuated by three major falls. There were peaks of origination in the Ludlow-Pragian, Givetian, late Famennian and Visean and peaks of extinction in the Pragian-Emsian, Givetian and early Tournaisian. Speciation and extinction rates were highest in the Lochkovian-Pragian and became progressively lower in subsequent stages. High correlation coefficients indicate that these events are associated with the availability of land habitat contingent on eustatic variations and increasing numbers of cosmopolitan genera. Meanwhile, proportions of endemic genera declined gradually. Due to less endemism and more migrations, both speciation and species extinction rates reduced. The changes of diversity and the timing of the three extinctions of land plants in South China are similar to those known already from Laurussia. However, the largest events in the Lochkovian-Pragian and subsequent smaller ones have not been seen in the global pattern of plant evolution. These land plant events do not correspond well temporally with those affecting land vertebrates or marine invertebrates. In South China, the diversity curve of land plants is generally opposite to that of marine faunas, showing a strong effect of eustatic variations. The increasing diversity of both land vertebrates and plants was punctuated above the Devonian-Carboniferous boundary, known as Romer's Gap, implying common underlying constraints on macroevolution of land animals and plants.

Earliest rooting system and root : shoot ratio from a new Zosterophyllum plant.

The enhanced chemical weathering by rooted vascular plants during the Silurian-Devonian period played a crucial role in altering global biogeochemical cycles and atmospheric environments; however, the documentation of early root morphology and physiology is scarce because the existing fossils are mostly incomplete. Here, we report an entire, uprooted specimen of a new Zosterophyllum Penhallow, named as Z. shengfengense, from the Early Devonian Xitun Formation (Lochkovian, c. 413 Myr old) of Yunnan, south China. This plant has the most ancient known record of a rooting system. The plant consists of aerial axes of 98 mm in height, showing a tufted habit, and a rhizome bearing a fibrous-like rooting system, c. 20 mm in length. The rhizome shows masses of branchings, which produce upwardly directed aerial axes and downwardly directed root-like axes. The completeness of Z. shengfengense made it possible to estimate the biomass allocation and root : shoot ratio. The root : shoot ratio of this early plant is estimated at a mean value of 0.028, and the root-like axes constitute only c. 3% of the total biomass. Zosterophyllum shengfengense was probably a semi-aquatic plant with efficient water use or a strong uptake capacity of the root-like axes.