Anatomical reinvestigation of Archaeopteris macilenta from the Upper Devonian (Frasnian) of South China

Abstract Archaeopteris macilenta is one of the most widespread plants in the Late Devonian. Based on fossils from the Frasnian Huangjiadeng Formation, Yichang District of Hubei Province, for the first time we study in detail the anatomy of this progymnosperm plant in South China. Ultimate axes are protostelic with three xylem sympodia and lack secondary tissue. Penultimate axes are eustelic, bearing eight sympodia and a thin band of secondary xylem. Radially symmetrical sympodia of mesarch primary xylem produce traces of appendages in a spiral arrangement. Archaeopteris macilenta and A. halliana (A. roemeriana) are dominant in the Frasnian and Famennian, respectively. Comparisons with these two species from other tectonic plates indicate consistent stelar architectures. Global spread, continuous occurrence, and identical anatomy during the Late Devonian indicate that Archaeopteris survived the Frasnian–Famennian extinction event. In this time, endemic genera and cosmopolitan taxa, including Archaeopteris, suggest the palaeogeographic isolation of South China and certain associations with other plates.     

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.     

Clevelandodendron ohioensis,gen. et sp. nov., a slender upright lycopsid from the late Devonian Cleveland Shale of Ohio

Clevelandodendron ohioensis Chitaley & Pigg gen. et sp. nov. is an almost entire lycopsid plant known from a single compressed specimen from the Cleveland Shale member of the Upper Devonian Ohio Shale. This unique specimen is 125 cm long, consisting of an unbranched, slender, monopodial axis with a partially preserved plant base bearing thick appendages at one end, and a compact, terminal ovoid bisporangiate strobilus at the other. The stem is 2 cm wide for most of its length. Visible on the decorticated stem surface are helically arranged, elongate leaf traces and laterally compressed, slender leaves along the stem margin. The plant base bears 4-6 thick appendages. The terminal strobilus is compact, ovoid, 9 cm long and up to 6 cm wide, morphologically similar to those of some Lepidodendrales, and bears helically arranged sporophyll/sporangium complexes with narrow bases and distal laminae up to 18 mm long, turned upward. Megaspores are 320-360 μm, trilete and laevigate, lacking a gula; microspores are 30-42 μm, trilete, indistinctly punctate and possibly assignable to Calamospora or Punctatisporites. Clevelandodendron demonstrates that slender unbranched lycopsids with an isoetalean plant habit similar to the Carboniferous genera Chaloneria and Sporangiostrobus and Triassic Pleuromeia-like forms were present as early as the Late Devonian. The early occurrence of this unique habit suggests that diversification within the isoetalean clade sensu Rothwell and Erwin (including both Isoetales and Lepidodendrales) was well established prior to the Carboniferous.     

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.     

ARCHAEOPTERIS MACILENTA,ANATOMY AND MORPHOLOGY OF ITS FROND

The stem, rachides, and pinnae of Archaeopteris macilenta, formerly considered to be a fern of Devonian age, comprise a branch system in which the ultimate divisions heretofore referred to as pinnules are the leaves. The primary vascular system of the “frond” is a lobed siphonostele from which leaf traces arise in a spiral sequence. The anatomy of the “rachis” and of the “pinnae” is shown to be similar to that of the stem, Callixylon, which bore these “fronds.” Branching, epidermal pattern and stomates are described for the spirally arranged leaves (fertile pinnules). Attachment and dehiscence of sporangia as well as their stomates are reported. Archaeopteris is retained in the Class Progymnospermopsida which includes plants with gymnospermous anatomy and pteridophytic reproduction. It is suggested that Actinopodium, Svalbardia and Siderella are related closely to Archaeopteris and that this group of genera shows evolutionary stages in webbing of leaves and planation of branch systems. The opportunities for ontogenetic studies of the arborescent genus Archaeopteris are pointed out.     

Devonian Vertebrates From Colombia

Vertebrate remains are reported from the Emsian–Eifelian Floresta Formation and the Late Devonian (?Frasnian) Cuche Formation of north‐eastern Colombia. The material from the Floresta Formation is associated with a marine invertebrate fauna and includes an arthrodire and probably a rhenanid. Several vertebrate‐bearing localities are recorded from the Cuche Formation; vertebrates occur with plant remains and lingulid fragments. They include an acanthodian (Cheiracanthoides? sp.), a chondrichthyan (Antarctilamna? sp.), placoderms (Bothriolepis sp., Asterolepis? sp. and an undetermined groenlandaspidid or primitive brachythoracid arthrodire), a stegotrachelid actinopterygianand three sarcopterygians (a cosmine‐covered form tentatively referred to an osteolepidid, the porolepiform Holoptychiussp., and the rhizodontid Strepsodus? sp.). This assemblage suggests a Late Frasnian age and is surprisingly similar to Late Devonian vertebrate assemblages found in similar facies of Europe and North America, notwithstanding the presence of the Gondwanan chondrichthyan Antarctilamna?. key words: Vertebrata, Devonian, Colombia, South America, biostratigraphy, palaeobiogeography.     

Microconchid tubeworms across the upper Frasnian – lower Famennian interval in the Central Devonian Field,Russia

Abstract: Microconchid tubeworms (Tentaculita) encrusting brachiopod shells have been investigated from the upper Frasnian – lower Famennian (Upper Devonian) deposits of the Central Devonian Field, Russia. The condition of microconchids and associated encrusting taxa is reported for the first time from the early Famennian recovery interval (crepida Chron) following the Frasnian–Famennian mass extinction. Two species, one new (Palaeoconchus variabilis sp. nov.) and the second one in open nomenclature (Palaeoconchus sp.), are described. Compared to lower Famennian specimens, they seem to be preferentially grouped on the anterior parts of the brachiopod host shells, which are interpreted as the most suitable sites away from the sea‐bottom and sediment. During the late Frasnian (Late rhenana Chron), microconchids, outnumbered by cornulitids and as abundant as foraminifers, were also associated with trepostome bryozoans, tabulates, rugose corals and various problematic encrusters. During the early Famennian recovery interval encompassing the crepida Chron, microconchids greatly outnumbered all associated encrusters, including the previously dominant cornulitids, while foraminifers, tabulates and rugose corals vanished. Early Famennian microconchids, represented by the single, albeit very abundant, species Palaeoconchus variabilis sp. nov., were opportunists that rapidly colonised the environment during the ongoing transgression following the regression‐driven biotic crisis in the area of the Central Devonian Field. In comparison to their late Frasnian predecessors and even other Middle Devonian specimens, no size reduction (the so‐called Lilliput effect) of early Famennian microconchid tubes was observed. It is probable that microconchids either rapidly attained their ‘normal’ sizes or they did not suffer any dwarfism following the Frasnian–Famennian event.     

TRILOBOXYLON ASHLANDICUM GEN. AND SP. N. FROM THE UPPER DEVONIAN OF NEW YORK

A new taxon is described from the Upper Devonian, Oneonta formation in the northern Cats-kills of New York. Triloboxylon gen. n. is represented by petrifactions showing two orders of branching. The main axis bears branches spirally and the latter bear the ultimate appendages spirally. Ultimate appendages branch dichotomously twice, in one plane. Primary xylem of the main axis and branches is three-armed with mesarch protoxylem extending in a continuous band within each arm. Primary xylem of the ultimate appendage is terete and dichotomizes twice. Metaxylem elements are characterized by scalariform and circular-bordered pitting on all walls. The cortex is composed entirely of isodiametrical parenchyma cells. Triloboxylon is compared with other genera with three-lobed protosteles. Its possible affinity with the Aneurophytales is shown. The morphological nature of the “frond” of the Aneurophytales and the possibility that the group possesses the morphological equivalent of simple leaves are discussed.     

Evolution of Devonian biodiversity of radiolarians with two porous spheres and one main spine

Features and major distinctions in the appearance and distribution of radiolarians with two porous spheres and one main spine in the Devonian basins are considered. Four solitary population waves and expansion scenario of radiolarians of this morphotype have been recognized in the Devonian. A new species, Radiobisphaera rozanovi sp. nov., from the Upper Emsian, Upper Eifelian, and Middle Frasnian of the southern Ural Mountains, from the Upper Eifelian of the Prague Basin (Barrandian), from the Middle-Upper Frasnian of the Rudnyi Altai, and from the Lower Famennian of the Timan-Pechora Basin and the Polar Ural Mountains is described.     

SVALBARDIA BANKSII SP. NOV. FROM THE UPPER DEVONIAN (FRASNIAN) OF NEW YORK STATE

Svalbardia banksii sp. nov., is described from the Upper Devonian (Frasnian) Fish Cabin Creek locality near Pond Eddy, New York. The compression consists of ultimate branches bearing spirally arranged, unwebbed leaves. The unwebbed leaves are up to 3.2 cm long, are somewhat flexuous, and dichotomize in more than one plane. The new species is similar to the reconstruction of Actinoxylon banksii from the Middle Devonian (Givetian) of New York and strengthens the supposed relationship between Svalbardia polymorpha and Actinopodium nathorstii from Mimerdalen, Spitzbergen.     

THE FROND OF HETERANGIUM

Frond members belonging to the monostelic seed fern genus Heterangium have been found in Pennsylvanian age coal balls collected in the Illinois Basin. Petioles bear small pinnae below a dichotomy which produced a bipartite frond. Pinnules of the Sphenopteris obtusiloba type are borne on secondary pinnae. The anatomy of each pinna order and the pinnules is described.     

PROBABLE ATTACHMENT OF THE DOLEROTHECA CAMPANULUM TO A MYELOXYLON-ALETHOPTERIS TYPE FROND

The discovery of pedunculate specimens of pollen organ Dolerotheca in close association with Myeloxylon-type pinnae and Alethopteris-type pinnules provides evidence for the reconstruction of a petrified medullosan frond and the attachment of the microsporangiate fructification. Specimens of Dolerotheca villosa and D. formosa from Middle and Upper Pennsylvanian age strata of North America are borne on slender peduncles with anatomical features and vascularization identical to those of some level in a Myeloxylon-Alethopteris-type frond. Reconstruction of the frond reveals a large, repeatedly dichotomising organ that bears penultimate pinnae and ultimate pinnules in a pinnate arrangement. Examination of pinnules on the surface of coal ball material indicates that they conform to the compression genus Alethopteris. Identical numbers and arrangement of vascular bundles together with identical anatomical features and multicellular hairs indicate that the Dolerotheca campanulum is borne in the position of a penultimate pinna. A reconstruction of the frond with a proposed polled organ attachment is included.     

New Late Devonian strophomenids (Brachiopoda) from Mongolia

Two new strophomenid species, Leptagonia barunkhuraica sp. nov. (Famennian) and Floweria mongolica sp. nov. (Frasnian), from the Upper Devonian of the Barunkhurai Depression of southern Mongolia are described.     

Environmentally controlled phyletic evolution, blindness and extinction in Late Devonian tropidocoryphine trilobites

The Middle and Upper Devonian carbonate succession of the Montagne Noire, Southern France has been precisely zoned by an unbroken sequence of conodont zones. Stratigraphic control is excellent, and has allowed evolutionary changes in tropidocoryphine trilobites, which occur throughout the succession, to be directly established. The tropidocoryphine had been a stable group for some 40 million years prior to the Middle Givetian. In their last few million years, however, they underwent rapid evolution and exhibit some striking transformations of the cephalon and the regression and virtual disappearance of the eye within a relatively short space of time. They also show a marked diminution in size, and lose their original relief so that the glabella becomes virtually flush with the surface. There are two separate lineages, both of which show eye-reduction and subsequent blindness. In the earlier lineage Tropidocoryphe (Longicoryphe)-Erbenicoryphe, the main features of the ancestral rootstock are conserved and the stable, strongly divergent anterior sutural pattern of the cephalon remains the same. The eye, however, became reduced to a slightly convex surface lacking lenses, only indistinctly defined. Erbenicorphe is confined to well-oxygenated facies, and probably lived as a shallow burrower within the sediment; it became extinct in the early Frasnian. The second lineage T. (Longicoryphe)-Pterocoryphe- Pteroparia shows a remarkable backward migration of the suture, which progressively swings posteriorly in successive species spanning four Frasnian conodont zones (about three million years duration). At the same time the eye progressively degenerates so that the last forms are blind. Sutural migration and eye reduction are not genetically linked, however; the unusual form of the cephalon and suture probably resulted from an adaptation to the euxinic environment in which Pterocoryphe originated. The loss of the eye resulted from the adoption of an endobenthic habit in Pteroparia which descended from the ancestral Pterocoryphe but which had migrated to an oxygenated facies. Eye-reduction is therefore parallel in the two lineages, but superimposed upon a different original cephalic configuration. The last Pteroparia became extinct when the late Frasnian fauna became overwhelmed by the first pulse of the ‘Kellwasser Event’ (probably an anoxic overturn). The evolving characters, through virtually all observed steps, show progressive unidirectional change without sudden breaks or saltations. Such unidirectional evolution is an adaptive response to constant long-lasting environmental influences. □Evolution, Devonian trilobites, France, gradualism, eye reduction.     

Anatomical reinvestigation ofArchaeopteris macilenta from the Upper Devonian (Frasnian) of South China

Archaeopteris macilenta is one of the most widespread plants in the Late Devonian.Based on fossils from the Frasnian Huangjiadeng Formation,Yichang District of Hubei Province,for the first time we study in detail the anatomy of this progymnosperm plant in South China.Ultimate axes are protostelic with three xylem sympodia and lack secondary tissue.Penultimate axes are eustelic,bearing eight sympodia and a thin band of secondary xylem.Radially symmetrical sympodia of mesarch primary xylem produce traces of appendages in a spiral arrangement.Archaeopteris macilenta and A.halliana (A.roemeriana) are dominant in the Frasnian and Famennian,respectively.Comparisons with these two species from other tectonic plates indicate consistent stelar architectures.Global spread,continuous occurrence,and identical anatomy during the Late Devonian indicate that Archaeopteris survived the Frasnian-Famennian extinction event.In this time,endemic genera and cosmopolitan taxa,including Archaeopteris,suggest the palaeogeographic isolation of South China and certain associations with other plates.     

Phytochrome control of rapid nyctinastic movements and membrane permeability in Albizzia julibrissin

Summary The rapid nyctinastic movements of Albizzia julibrissin pinnules are under the control of phytochrome. When given prior to a dark period, red light facilitates and far-red light inhibits pinnule closure in the dark. These light effects are mutually photoreversible. The opening reaction of the pinnules following a dark period is mediated mainly by blue light. The nyctinastic closure response is accompanied by an increased rate of electrolyte efflux from the cut pinna base. This observation, coupled with the fact that the rapid nyctinastic movement is not affected by actinomycin D, supports the view that phytochrome control of the sleep movement is not mediated through effects on RNA metabolism, but rather through changes in membrane permeability.     

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.     

Devonian Gastropoda from the Dornap “Massenkalk” complex (Bergisches Land,Germany)

Seven Middle Devonian (latest Givetian or earliest Frasnian) gastropod species are described from the quarry of Oetelshofen near Dornap (Bergisches Land, northwestern Rheinisches Schiefergebirge, Germany). New species areMurchisonia dornapica, Araeonema osterholzensis, Plagiothyra isekei andMacrochilina drozdzewski. Additionally, a new subspecies,Euryzone delphinuloides oetelshofensis and aNaticopsis-like species and their typical facial occurrence are discussed. The existence of predation is proved by a bore-hole inMacrochilina drozdzewski; the predator itself is not yet identified with certainty.      

The fate of the homoctenids (Tentaculitoidea) during the Frasnian–Famennian mass extinction (Late Devonian)

The homoctenids (Tentaculitoidea) are small, conical‐shelled marine animals that are among the most abundant and widespread of all Late Devonian fossils. They were a principal casualty of the Frasnian–Famennian (F‐F, Late Devonian) mass extinction, and thus provide an insight into the extinction dynamics. Despite their abundance during the Late Devonian, they have been largely neglected by extinction studies. A number of Frasnian–Famennian boundary sections have been studied, in Poland, Germany, France, and the USA. These sections have yielded homoctenids, which allow precise recognition of the timing of the mass extinction. It is clear that the homoctenids almost disappear from the fossil record during the latest Frasnian ‘Upper Kellwasser Event’. The coincident extinction of this pelagic group, and the widespread development of intense marine anoxia within the water column, provides a causal link between anoxia and the F‐F extinction. Most notable is the sudden demise of a group, which had been present in rock‐forming densities, during this anoxic event. One new species, belonging to Homoctenus is described, but is not formally named here.