The youngest Ediacaran fossils from southern Africa

Discovery of fossils of the Ediacara biota near the top of the Spitzkopf Member at farm Swanpunt extends the known range of these remains in Namibia more than 600 m to near the sub-Cambrian unconformity. The fossiliferous beds occur approximately 100 m above a volcanic ash dated at 543 +/- 1 Ma, and thus may be the youngest Proterozoic Ediacara-type fossils reported anywhere in the world. Fossils are preserved within and on the tops of dm-thick beds of storm-deposited sandstone at two stratigraphic levels; the environment is interpreted as open marine, generally calm but with episodic disruptions by storm waves, and probably within the euphotic zone. The presence of Pteridinium carolinaense (St. Jean), which is also known from the classic sections in Ediacara and the White Sea among others, reinforces evidence from geochronology and chemostratigraphy that the Swartpunt section is terminal Neoproterozoic in age. The new genus and species Swartpuntia germsi is a large, multifoliate frond that exhibits at least three quilted petaloids. Macroscopically, Swartpuntia resembles Pteridinium and Ediacara-type fronds such as Charniodiscus traditionally interpreted as Cnidaria, whereas microscopically it exhibits segmentation that is remarkably similar to that of the putative worm Dickinsonia. Combination of diagnostic characters of these supposedly disparate groups in a single species suggests that many species of quilted Ediacaran organisms were more similar to each other than they were to any modern groups, and provides support for the concept of the "Vendobionta" as a late Neoproterozoic group of mainly multifoliate organisms with a distinctive quilted segmentation.     

Anatomical and ontogenetic reassessment of the Ediacaran frond Arborea arborea and its placement within total group Eumetazoa

Organisms in possession of a frondose body plan are amongst the oldest and most enigmatic members of the soft‐bodied Ediacaran macrobiota. Appraisal of specimens from the late Ediacaran Ediacara Member of South Australia reveals that the frondose taxon Arborea arborea probably possessed a fluid‐filled holdfast disc, the size and form of which could vary within populations. Mouldic preservation of internal anatomical features provides evidence for tissue differentiation, and for bundles of tubular structures within the stalk of the organism. These structures connect in a fascicled arrangement to individual lateral branches, before dividing further into individual units housed on those branches. The observed fascicled branching arrangement, which seemingly connects individual units to the main body of the organism, is consistent with a biologically modular construction for Arborea, and raises the possibility of a colonial organization. In conjunction with morphological characters previously recognized by other authors, including apical‐basal and front‐back differentiation, we propose that to the exclusion of all alternative known possibilities, Arborea can be resolved as a total group eumetazoan.     

Testing the protozoan hypothesis for Ediacaran fossils: a developmental analysis of Palaeopascichnus

Abstract: The hypothesis that the Ediacara biota were giant protozoans is tested by considering the external morphology, internal organization, suggested fossil representatives and molecular phylogeny of the xenophyophores. From this analysis, we find no case to support a direct relationship. Rather, the xenophyophores are here regarded as a group of recently evolved Foraminifera and are hence unlikely to have a record from the Ediacaran Period. Further from the growth dynamics of Foraminifera, they are also unlikely to be related to the Palaeopascichnus organism. We also find significant distinctions in the growth dynamics of Palaeopascichnus and organisms usually referred to the Ediacara biota, such as Charnia and Dickinsonia. Developmental analysis of the Palaeopascichnus– central to the xenophyophore hypothesis – reveals unusual, protozoan features, including evidence for chaotic repair structures, for mergence of coeval forms, as well as complex bifurcations. These observations suggest that Palaeopascichnus is a body fossil of an unidentified protozoan but is unrepresentative of Ediacaran body construction, in general.     

ABSCISSION AND THE AXILLARY FROND IN SPIRODELA OLIGORHIZA (LEMNACEAE)

Two abscission zones are present in fronds of Spirodela oligorhiza. In one zone the abscission layer separates the daughter frond and its connecting stalk from the mother frond. The second abscission layer separates the daughter frond from the connecting stalk. An axillary frond is asymmetrically positioned at the base of each daughter frond where it joins the mother frond. In right-handed physiological clones the axillary frond is to the right of the connecting stalk and in left-handed clones, to the left of the stalk. When a daughter frond abscises it leaves behind its axillary frond in the pocket of the mother frond. Histological features of abscission and treatments that induce abscission in Spirodela oligorhiza are described.     

湖北三峡地区埃迪卡拉系灯影组Hiemalora stellaris生物属性及埋藏分析

Hiemalora stellaris是湖北宜昌三峡地区埃迪卡拉系灯影组石板滩生物群中数量最为丰富的化石类型之一,由位于中心位置的圆盘和围绕盘缘向外伸出的似触手状结构组成,似触手状结构长度与圆盘直径具有较好相关性。同时也发现少量Hiemalora与茎干相连,其中一枚产自石板滩段底部的标本保存完整,与茎干连接有叶状体。叶状体呈卵形,可见一级分枝与二级分枝,顶部有呈"牛角状"的尖刺,叶状体形态与Charniodiscus spinosus相似。结合纽芬兰地区发现的Primocandelabrum hiemaloranum,表明Hiemalora作为生物体的固着器官,起到抵御水流冲刷、支撑和固定整个生物体的作用,但与其相连的叶状体表现出不同的形态类型,表明Hiemalora化石可能具有较为多样的生物来源。根据化石的形态特征和埋藏状态分析,Hiemalora在生物生活时位于藻席层之下的沉积物内,常呈正凸起保存于岩层顶面,而茎干和叶状体位于水体中,生物死亡后,叶状体倒伏在藻席层之上,当岩层沿藻席层劈开后,在顶面保存为凹陷的印痕。     

Twin shoots borne on a Botryopteris frond from the British upper carboniferous

A long rachis of Bottyopteris aff. hirsuta bearing twin opposite lateral shoots is described from a coal ball of Westphalian age collected in Lancashire. The simultaneous emission, in the middle region of a frond, of two short axes each bearing a large number of poorly developed leaves is a feature hitherto unrecorded for the genus. The specimen suggests that this Paleozoic fern consisted of long fronds whose branching pattern showed a certain capacity for morphological variation; it provides yet another example of the surprisingly limited development of shoots borne on Botryopteris fronds.     

Sind Farne Kormophyten? Eine Alternative zur Telomtheorie

Modern ideas concerning cormophyte phylogeny are strongly influenced by the telome theory. Erect growth, radial symmetry, dichotomy, protostely and eusporangial structure are considered to be primitive features. Yet, the cormus needs redefinition. TheHypolepidaceae (s. str.) are shown to have rather a two-dimensional thallus than a three-dimensional cormus. Their so-called rhizome develops a continuous areophore margin which is connected with the margin of the fronds. The longitudinally inserted fronds and branches are produced by the terminal rhizome meristem which itself grows in many respects like the apical frond meristem. The rhizomes ofDennstaedtiaceae and theDavallia-type of rhizomes are intermediate between thisHypolepis-type and the true cormus-type. TheHypolepis-type is compared with fern prothalli. Under perhumid tropical conditions higher land plants may have evolved from small, creeping, two-dimensional fern prothallium-like progenitors with isomorphous gameto- and sporophytes.

Vorgetragen auf der Morphologentagung in Zürich 1975.     

Eggs Distribution Behaviour of Dubas Bug (Ommatissus Lybicus: Homoptera: Tropiduchidae) in Relation to Seasons and Some Physico- Morphic Characters of Date Palm Leaves

Dubas bug, Ommatissus lybicus, is a major pest of date palm in Panjgur Balochistan. Both the nymphs and adults suck the sap from fruit stalk, leaves and may cause about 50 % of economic losses. Chlorophyll contents and leaf thickness was measured by using SPAD-502 chlorophyll meter and vernier caliper respectively. Eggs distribution density was highly significant in different frond rows and within frond during spring (1^st) and summer (2^nd) generations. In both generations number of eggs per leaflet decreased towards the apex fronds rows and within frond number of eggs decreased towards the terminal leaflets. Eggs distribution had negative correlation with leaf thickness; chlorophyll contents (SPAD value) and frond row (bottom to top). Apparently shade affect the eggs laying behaviour of dubas bug.     

Translocation of C in Macrocystis pyrifera (Giant Kelp)

The pattern of import and export of ¹⁴C-labeled assimilates in Macrocystis pyrifera (L.) C. A. Agardh in southern California was studied by labeling single blades on fronds, in situ, with [¹⁴C]NaHCO₃ for 24 hours. The pattern was found to be similar to that known in dicotyledons: actively growing tissue imported and did not export. As a blade reached maturity it began to export, at first only acropetally to the apex which formed it, later also down the frond to sporophylls and frond initials at the base of the frond, and into the apical regions of juvenile fronds; finally there was a phase of declining export, late in the life of the blade, when transport was only downward. Young fronds imported from older fronds until they were approximately 3 meters long, by which time they had developed mature, upward exporting blades. No translocation was found from a younger frond to an older frond, nor was there transport upward from a blade on a frond lacking the apical region.     

Microbial biofilms and the preservation of the Ediacara biota

Laflamme, M., Schiffbauer, J.D., Narbonne, G.M., & Briggs, D.E.G. 2011: Microbial biofilms and the preservation of the Ediacara biota. Lethaia, Vol. 44, pp. 203–213. The terminal Neoproterozoic Ediacaran Period is typified by the Ediacara biota (ca. 579–542 Ma), which includes the first morphologically complex macroscopic organisms. Both the taphonomic setting that promoted the preservation of the soft‐bodied Ediacara biota in coarse‐grained sediments, and the influence of associated microbial coatings on this process, have generated debate. Specimens of Ediacaran discs (Aspidella) from the Fermeuse Formation of Newfoundland, Canada, were analysed using environmental scanning electron microscopy (ESEM) and focused ion beam electron microscopy (FIB‐EM) to determine the relationship between the fossil specimens and the surrounding sediment. The presence of chemically distinct (Al–Mg–Fe–K‐ and to a lesser extent S‐rich), finer‐grained sediment (with organized iron sulphides) surrounding the upper and lower margins of the Ediacaran fossils is consistent with elemental analyses of well preserved bacterial biofilms from other localities. ESEM analyses reveal a contrast in the composition of the sediment bound within the discs, which contains a higher concentration of Al, Ca and K, and the purer Si‐rich sediment that forms the surrounding matrix. This suggests that the coarse grained sediment was incorporated into the organism during life. Ediacaran discs were likely surrounded by a bacterial biofilm or thin microbial mat composed primarily of extracellular polymeric substances (or exopolysaccharide) during life, which added structural stability to these frond holdfasts, and facilitated their fossilization. Microbially mediated preservation in Fermeuse‐style Ediacaran taphonomy provides an explanation for the dominance of Aspidella holdfasts in these settings, and suggests that preservation of Ediacaran fossils in the round may be much more prevalent than previously recognized. We suggest that the overwhelming dominance of circular to bulbous forms such as Aspidella in Ediacaran biotas around the world is a direct result of the interplay between microbial ecology and microbially mediated taphonomy. □Aspidella, Ediacaran preservation, environmental scanning electron microscopy, focused ion beam electron microscopy, palaeoecology, taphonomic bias.     

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.     

Effaced preservation in the Ediacara biota and its implications for the early macrofossil record

Abstract: Ediacaran structures known as ‘pizza discs’ or Ivesheadia have long been considered enigmatic. They are amongst the oldest known members of the Ediacara biota, apparently restricted to the Avalonian successions of Newfoundland and the UK, c. 579–560 Ma. Here, we suggest that these impressions are taphomorphs, resulting from the post‐mortem decay of the frondose Ediacaran biota. Ediacaran fossils range from well‐preserved, high‐fidelity variants to almost completely effaced specimens. The effaced specimens are inferred to have undergone modification of their original morphology by post‐mortem microbial decay on the sea floor, combined with sediment trapping and binding. In this style of preservation, morphological details within the organism became variously subdued as a function of the extent of organic decay prior to casting by overlying sediments. Decay and effacement were progressive in nature, producing a continuum of grades of preservation on Ediacaran bedding planes. Fossils preserved by such ‘effaced preservation’ are those that have suffered these processes to the extent that only their gross form can be determined. We suggest that the lack of detailed morphology in effaced specimens renders such fossils unsuitable for use as type material, as it is possible that several taxa may, upon degradation and burial, generate similar morphological taphomorphs. We here reinterpret the genus Ivesheadia as a taphomorph resulting from extensive post‐mortem decay of frondose organisms. Blackbrookia, Pseudovendia and Shepshedia from beds of comparable age in England are likewise regarded as taphomorphs broadly related to Charnia or Charniodiscus spp. To reflect the suggestion that such impressions are likely to be taphomorphs, and not taxonomically discrete, we propose the term ivesheadiomorphs to incorporate all such effaced taphonomic expressions of Ediacaran macrofossil taxa in Avalonian assemblages. Our recognition of effaced preservation has significant implications for Ediacaran taxonomy, and consequently for measures of Ediacaran diversity and disparity. It is implied that Avalonian assemblages preserve both organisms that were alive and organisms that were already dead at the time of burial. As such, the fossil assemblages cannot be taken to represent census populations of living organisms, as in prior interpretations.     

Plant response to overcrowding – Lemna minor example

Plants adopt various strategies in response to increasing density. We tested that response in two populations of Lemna minor L. – a free floating aquatic plant that frequently experiences intraspecific competition for space. Surface area of fronds and colonies, colony size (the number of fronds per colony), the rate of reproduction (based on the number of produced fronds) and growth rate (enlargement of surface area of all colonies) were the analysed factors presumably affected by density. The study was performed in natural stands and in experimental conditions with the use of two contrasting plant densities. Plants growing in natural conditions produced fronds of smaller and less variable surface area as a response to overcrowding but the number of fronds per colony was unrelated to plant density. Stable experimental conditions facilitated formation of fronds and colonies larger than in the field but frond detachment decreasing colony size was more intensive at high than at low density. This strategy allowed plants to more efficiently occupy limited available space. No self-thinning was observed during experimental cultures. Due to increasing frond area in cultures, growth rate was always higher than the rate of plant reproduction. Both were strongly negatively affected by high density. Performed calculations indicate that density-dependent growth inhibition starts when L. minor colonies cover the available water surface with a mono-layer mat. Some types of responses were found to significantly differ between analysed populations, which was also shown by genetic differences tested with he ISSR-PCR technique. Possible causal relationship between plant strategies and their genomic structure needs, however, further studies.     

The Structure and Development of the Thallus in the British Species of Gelidium and Pterocladia

The apical structure and the development of the thalli of allthe British species of Gelidium and Pterocladia have been investigated;the development of G. pulchellum is described in detail. Eachaxis is terminated by one or more apical cells, which by theirsegmentation form the tissues of the thallus. An axial filamentis distinct for a short distance behind each apical cell, butsecondary pit-connexions develop rapidly so that in sectionthe mature axis has the appearance of a multi-axial structure. Lateral branches of the frond develop by the segmentation ofthe lateral branch apical cells, which are formed by the transformationof superficial cortical cells, either in the meristematic ormature parts of the axes. The extreme variability of externalappearance is due principally to the indeterminate origin ofall lateral branches. The thallus in the British species of Gelidium and Pterocladiaconsists of erect fronds borne on creeping axes. The relativeproportions of the frond and creeping axes in various speciesand their survival through adverse conditions are discussed.     

Reassessment of the Devonian problematicum Protonympha as another post‐Ediacaran vendobiont

Protonympha is an enigmatic fossil represented by two species from the Middle Devonian (Protonympha transversa) and Late Devonian (Protonympha salicifolia) of New York. Although interpreted in the past as a polychaete worm or starfish arm, Protonympha is not found with marine fossils, but with fossil plants. This fossil plant community was a swamp woodland of Lepidosigillaria whitei, with ground cover of Haskinsia colophylla, fringing brackish to freshwater coastal lagoons of the Catskill Delta. Protonympha shares with Ediacaran Vendobionta a quilted body of unskeletonized biopolymer that is unusually resistant to burial compaction. In overall form, Protonympha is most like the Ediacaran genus Spriggina. Protonympha has branching and tapering tubular structures radiating from the bottom. These rhizine‐like structures, thallus stratification and internal chambers revealed by petrographic thin sections suggest affinities with lichenized fungi. As for Cambrian Swartpuntia and Ordovician–Silurian Rutgersella, Protonympha may have been a post‐Ediacaran vendobiont.     

Fecundity,spore recruitment and size in Gelidium sesquipedale (Gelidiales,Rhodophyta)

Gelidium sesquipedale fecundity was quantified by counting tetrasporangial sori and cystocarps per meter squared and by estimating the number of spores contained inside them. These were obtained by regression on a size metric of reproductive structures. Tetrasporangial sori length and cystocarp thickness were the best estimators of spore number. To assess spore recruitment, 12 pottery tiles were fixed to the bottom, and the appearance of small fronds was monitored.No clear seasonal pattern of reproduction was found. Tetraspore production peaked in March 1990 with 10.4 × 10⁶ spores m^–2, whereas the carpospore peak was lower, 4.9 × 10⁵ spores m^–2 in July 1989. Recruitment followed tetraspore peaks. The probability of a G. sesquipedale tetraspore making the transition to a recruit was 4.7 × 10^–5. Frond length was significantly related to tetrasporangial sori number, while cystocarp number was only related to frond branching order. Minimum size for reproduction was 6.9 cm for gametophytes and 5.4 cm for tetrasporophytes; very rarely were cystocarpic fronds smaller than 9 cm, while tetrasporic fronds were often longer than 15 cm. Cystocarpic fronds were significantly shorter and had more branches than tetrasporic fronds.     

TRANSLOCATION OF 14C IN MACROCYSTIS INTEGRIFOLIA (PHAEOPHYCEAE)1,2

Translocation patterns in the giant kelp, Macrocystis integrifolia Bory, were investigated in situ using ¹⁴C tracer; sources and sinks were identified. Export was first detected after 4 h of labeling; experiments were routinely 24 h continuous ¹⁴C application. Mature blades exported ¹⁴C to young blades on the same frond and on younger fronds, as well as to sporophylls and frond initials at the bases of the fronds. Blades <0.3 m from the apex imported and did not export; this distance did not change seasonally. In spring export from blades 0.3–1.25 m from the apex was exclusively upwards; older blades also exported downwards. In fall downward export began 0.5 m from the apex, and blades >2 m from the apex exported exclusively downwards. Carbon imported by frond initials, young fronds, and sporophylls in fall may partly be stored for growth in early spring. No translocation was seen in very young plants until one blade (secondary frond initial) bad been freed from the apical blade; this blade exported to the apical blade for a time, but imported when it began to develop into a frond. The second and third formed blades on the primary fronds (sporophylls also exported when <0.3 m from the apex, and later stopped. Frond initials and sporophylls on later-formed fronds did not export at all. The translocation pattern in M. integrifolia differs from that previously reported in M. pyrifera in seasonal change and in distances from the apex at which the changes take place.     

Global marine redox changes drove the rise and fall of the Ediacara biota

The role of O₂ in the evolution of early animals, as represented by some members of the Ediacara biota, has been heavily debated because current geochemical evidence paints a conflicting picture regarding global marine O₂ levels during key intervals of the rise and fall of the Ediacara biota. Fossil evidence indicates that the diversification the Ediacara biota occurred during or shortly after the Ediacaran Shuram negative C‐isotope Excursion (SE), which is often interpreted to reflect ocean oxygenation. However, there is conflicting evidence regarding ocean oxygen levels during the SE and the middle Ediacaran Period. To help resolve this debate, we examined U isotope variations (δ²³⁸U) in three carbonate sections from South China, Siberia, and USA that record the SE. The δ²³⁸U data from all three sections are in excellent agreement and reveal the largest positive shift in δ²³⁸U ever reported in the geologic record (from ~ ?0.74‰ to ~ ?0.26‰). Quantitative modeling of these data suggests that the global ocean switched from a largely anoxic state (26%–100% of the seafloor overlain by anoxic waters) to near‐modern levels of ocean oxygenation during the SE. This episode of ocean oxygenation is broadly coincident with the rise of the Ediacara biota. Following this initial radiation, the Ediacara biota persisted until the terminal Ediacaran period, when recently published U isotope data indicate a return to more widespread ocean anoxia. Taken together, it appears that global marine redox changes drove the rise and fall of the Ediacara biota.