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.

A Late Devonian tree lycopsid with large strobili and isotomous roots.

Tree lycopsids prospered in the Late Devonian and constituted a major part of the Late Paleozoic forest ecosystem that deeply impacted the Earth's climate. However, the fertile organs of these early tree lycopsids display low morphological disparity, which has hampered further knowledge about their ecological habit. Here, we report Omprelostrobus gigas gen. et sp. nov. from the Upper Devonian (Famennian) Wutong Formation at Changxing, Zhejiang, China. The collection includes aerial axes, strobili and associated roots. The strobili are the largest among coeval lycopsids to our knowledge, and are divided into proximal and distal portions by dimorphic sporophylls with differentiated laminae and probable strong photosynthetic capacity. The associated but not attached roots displaying multiple isotomous branches lack rootlets and typical rootlet scars. The varied strobili sizes of early tree lycopsids were relatively independent of their body plan, but the large strobili could suggest increased reproductive investment to overcome the disadvantages of the disturbed flooded habitat.

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.

Pollen Organ Telangiopsis sp. of Late Devonian Seed Plant and Associated Vegetative Frond.

Pollen organ Telangiopsis sp., associated with but not attached to vegetative fronds, has been collected from the Upper Devonian (Famennian) Wutong Formation, Dongzhi County, Anhui Province, China. Fertile axes with terminal pollen organs are dichotomous for 2-4 times and may be proximally attached by fragmentary pinnules. Pollen organs are synangiate and borne on the top of a short stalk. Synangia are radial in symmetry and each consists of 4-8 elongate microsporangia fused at base. Microsporangia have a longitudinal dehiscence line and show a tapered apex. The associated stem is spiny and bears a vegetative frond which bifurcates once at the basalmost part. Frond rachises possess one order of pinna arranged alternately. Pinnules are borne alternately, planate, highly dissected, and equally dichotomous for 2-3 times. Comparisons among Late Devonian seed plants recognize several branching patterns in the fertile fronds/axes bearing terminal pollen organs. Telangiopsis sp. reinforces that the Late Devonian pollen organs are synangiate usually with basally fused microsporangia. It is suggested that the evolutionary divergence of radial and bilateral symmetries of pollen organs may have occurred in the Famennian, when the earliest seed plants evolved planate and sometimes laminate pinnules.

Vegetative characters, growth habit and microsporangiate strobilus of lycopsid Minostrobus chaohuensis.

Late Devonian Minostrobus chaohuensis is one of the earliest monosporangiate-strobilate isoetaleans. Based on new material of this plant, the vegetative axis and microsporangiate strobilus are studied in detail, and the whole plant knowledge is summarized. The vegetative axis is isotomously branched. The stem is up to 55 mm in diameter with helically arranged leaf cushions. Stems and thick branches bear long fusiform leaf cushions and interareas with vertical linear ornamentations. A ligule pit, oblanceolate leaf scar, and vascular bundle scar appear on the leaf cushion. Distal axes have persistent lanceolate leaves and rhombic leaf bases. The microsporangiate strobilus is cylindrical in shape, possesses sporophyll with alate pedicel and long triangular lamina, uniseriate sporangial wall, subarchesporial pad inside the sporangium, and microspore with cingulum. Based on comparisons with other isoetaleans, the usage of the terms "leaf cushion" and "leaf base" is discussed, and Minostrobus chaohuensis is considered as a tree-like lycopsid. It suggests that arborescent isoetaleans with monosporangiate strobili had appeared and diversified in the Late Devonian. The multi-dichotomous branching system of Minostrobus provides new data on the evolution of growth architecture in rhizomorphic lycopsids.