| 1. |
- Agić, Heda, 1989-, et al.
(författare)
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Affnity, life cycle, and intracellular complexity of organic-walled microfossils from the Mesoproterozoic of Shanxi, China
- 2015
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Ingår i: Journal of Paleontology. - : Cambridge Journals. - 0022-3360 .- 1937-2337. ; 89:1, s. 28-50
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Tidskriftsartikel (refereegranskat)abstract
- Light microscope and scanning electron microscope observations on new material of unicellularmicrofossils Dictyosphaera macroreticulata and Shuiyousphaeridium macroreticulatum, from the MesoproterozoicRuyang Group in China, provide insights into the microorganisms’ biological affinity, life cycle and cellularcomplexity. Gigantosphaeridium fibratum n. gen. et sp., is described and is one of the largest Mesoproterozoicmicrofossils recorded. Phenotypic characters of vesicle ornamentation and excystment structures, properties ofresistance and cell wall structure in Dictyosphaera and Shuiyousphaeridium are all diagnostic of microalgalcysts. The wide size ranges of the various morphotypes indicate growth phases compatible with the development ofreproductive cysts. Conspecific biologically, each morphotype represents an asexual (resting cyst) or sexual (zygotic cyst)stage in the life cycle, respectively. We reconstruct this hypothetical life cycle and infer that the organism demonstrates areproductive strategy of alternation of heteromorphic generations. Similarly in Gigantosphaeridium, a metabolicallyexpensive vesicle with processes suggests its protective role as a zygotic cyst. In combination with all these charactersand from the resemblance to extant green algae, we propose the placement of these ancient microorganisms in the stemgroup of Chloroplastida (Viridiplantae). A cell wall composed of primary and secondary layers in Dictyosphaera andShuiyouisphaeridium required a high cellular complexity for their synthesis and the presence of an endomembranesystem and the Golgi apparatus. The plastid was also present, accepting the organism was photosynthetic. The biotareveals a high degree of morphological and cell structural complexity, and provides an insight into ongoing eukaryoticevolution and the development of complex life cycles with sexual reproduction by 1200Ma.
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| 2. |
- Agić, Heda, 1989-, et al.
(författare)
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Cyst and operculum formation in Cambrian-Ordovician galeate acritarchs from Estonia : implications for the algal phylogeny and blooms in the early Paleozoic
- 2014
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Ingår i: 4th International Palaeontological Congress Abstract Volume. ; , s. 913-
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Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract
- Unicellular organic-walled microfossils have been recovered from the Cambrian Lükati Formation and the Tremadocian Varangu Formation exposed in northern Estonia. Due to a combination of main morphological and biochemical characters, mainly a) excystment opening, b) processes, c) acetolysis- resistant vesicle wall, microfossils have been interpreted as reproductive cysts of green algae. Both microfossil assemblages reflect the evolutionary patterns though the early Palaeozoic: from the Cambrian radiation of morphologically innovative taxa to increase in diversity and more disparate Ordovician forms. Combined light transmitted and scanning electron microscopy on the Middle Cambrian to Tremadocian galeate plexus acritarchs Caldariola, Priscogalea and Stelliferidium, revealed exceptionally preserved morphological elements and rare structure among fossil and extant microbiota – an opening with operculum (lid) in reproductive cyst, in addition to lavish vesicle ornamentation and sculpture. Operculum formation model is reconstructed from fossils at different stages of operculum position and attachment. Comparative morphology shows strong similarity of galeates to the reproductive cysts of the extant algae of Dasycladales (Chlorophyta), where the lid covering the cyst opening is determined by an intrinsic lid-forming apparatus during the organism’s reproductive stage. Opercula in Cambro-Ordovician galeate acritarchs and Dasycladales may be considered a homologous character. Unique morphology of the operculum-bearing microbiota would have required a degree of intracellular sophistication for its development, suggesting advanced intracellular machinery present already in the early Palaeozoic phytoplankton. Additionally, a new species of minute, sphaeromorphic and aggregated eukaryotic microfossils is recorded. It possesses a vesicle wall with corrugated sculpture and perforated by nano-scale pores. These minute early Cambrian microfossils have diagnostic characters of prasinophyte algae.
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| 3. |
- Agić, Heda, 1989-, et al.
(författare)
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Diversity of organic-walled microfossils from the early Mesoproterozoic Ruyang Group, North China Craton - a window into the early eukaryote evolution
- 2017
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Ingår i: Precambrian Research. - : Elsevier BV. - 0301-9268 .- 1872-7433. ; , s. 101-130
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Tidskriftsartikel (refereegranskat)abstract
- Mesoproterozoic Era was an important time for the initial diversification of eukaryotic groups and the appearance of the first complex morphologies. While eukaryotes evolved around 2.4 Ga, the first microfossils with ornamentation and sculpture occur in the 1.8-1.6 Ga successions worldwide. Shales and siltstones of the Ruyang Group, Shanxi Province, North China Craton, record a high diversity of such organic-walled microfossils. Recently, the depositional ages of this succession has been constrained to 1.75-1.40 Ga via zircon U-Pb dating. This dating extends back the time of the first appearance of complex eukaryotic characters (e.g. processes, complex wall structure) in the fossil record. We have conducted a biostratigraphic investigation on of the samples throughout the fossiliferous Ruyang Group to provide an estimate of the early eukaryotic diversity in the Mesoproterozoic. Light- and scanning electron microscope studies have documented 26 species, including several that are reported for the first time, and some that were previously known only from younger, Neoproterozoic strata. Fossil diversity is high in the upper Baicaoping Formation, declines in the middle and reaches its peak in the upper Beidajian Formation. Novel morphologies among the unicellular Ruyang biota include a variety of processes, from tube-like extensions to hirsute spines, vesicles with velutinous outer membranes, as well as numerous specimens with internal bodies of varying sizes. We have also recorded the globally distributed Mesoproterozoic taxa Dictyosphaera, Shuiyousphaeridium, and Tappania. Key characters displayed by the Ruyang biota are consistent with reproductive structures (especially cysts among modern protists. These microfossils provide an additional evidence for the emergence of the crown group Eukarya by 1.7-1.4 Ga.
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| 4. |
- Agić, Heda, 1989-, et al.
(författare)
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Is cyst formation in early eukaryotes a requirement for their preservation in the fossil record?
- 2015
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Ingår i: Abstracts of the Astrobiology Science Conference 2015: Habitability, Habitable Worlds and Life.
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Konferensbidrag (refereegranskat)abstract
- Most of the Archaean-Proterozoic fossil record consists of non-biomineralizing microorganisms or their signatures. Body fossils of bacteria and early eukaryotes are preserved in siliciclastics, shales and carbonates, and are usually studied by preparation of thin sections or extraction from the rock matrix via acid maceration.The first eukaryotic organic-walled microfossils (OWM) appear at least by 1.8 Ga and undergo morphological diversification and evolutionary radiation in the Mesoproterozoic. There are no preserved eukaryotic-grade microfossils except OWM until the onset of biomineralization much later in the Neoproterozoic, evident in the record of testate amoebae (VSM) and microfossils with scaly elements.OWM are a less conspicuous component of the fos-sil record than taxa with skeletal or shelly elements. Organic matter decays quickly upon death of the organism, due to autolytic enzymes or degradation via het-erotrophy. However, species producing vegetative cells, resting cysts, zygotes, or spores, show considerable resistance to autolysis. Case studies on extractable carbonaceous OWM indicate they are preserved due to complex refractory molecules in the structure of their sturdy vesicle walls. Living analogues across protistan clades utilise such sporopollenin-like compounds for the cyst wall construction during reproductive phase. Algaenan-containing trilaminar sheath structure (TLS) is secreted during aplanospore formation in extant chlorophyte alga Haematococcus. TLS has also been documented in Leiosphaeridia acritarchs from the Cambrian Lükati Formation in Estonia. Leiosphaeridia is a long ranging morphotype, dating as far back as 1.8 Ga. Presence of TLS in these fossils suggests their function as reproductive cyst. Dictyosphaera-Shuiyousphaeridium plexus from the Mesoproterozoic Ruyang Group, China, also exhibits cyst-like morphology and unique elements of wall reinforcement: internally secreted organic platelets.In addition to these early OWM, many Meso-Neoproterozoic taxa such as Tappania, Trachyhystrychosphaera and Kildinella contain cyst-like characters: 1) reproductive openings, 2) ornamentation, 3) occa-sionally preserved internal bodies and 4) acetolysis-resistant vesicle walls – properties observed among extant encysting protists.Ornamented (process-bearing) microfossils in par-ticular bear strong similarities with zygotes of living unicellular algae. Property of acetolysis-resistant vesicle is a result of sporopollenin-like macromolecules in the wall, synthesized most commonly by the autotrophic eukaryotes. Presence of such recalcitrant organic walls requires significant metabolic investment by the microorganism, which suggests a protective and/or reproductive function. This also allows for easier, and more detailed preservation in the rock record.One of the concerns arising from the studies on the early eukaryotic fossils is the bias towards encysting organisms. The eventual search for the fossil record on other planetary bodies could face the same challenges as the Precambrian palaeobiology: fossilisation and eventual detection might be problematic for any unicellular eukaryotic-grade organisms if they have not evolved reproduction via encystment, or cyst formation as means of coping with adverse environmental conditions.
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| 5. |
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| 6. |
- Agić, Heda, 1989-, et al.
(författare)
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Morphology of the Proterozoic eukaryotic microfossils as a reflection of their intracellular complexity
- 2014
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Ingår i: 4th International Palaeontological Congress Abstract Volume. ; , s. 256-
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Konferensbidrag (refereegranskat)abstract
- Mesoproterozoic is a time of increasing diversity of microscopic life and appearance of intricate new cell morphologies. First eukaryotes may have evolved around 2.4 Ga, but the first microbiota with intricate sculpture and ornamentation are found in the younger, 1.8.-1.6 Ga successions worldwide. Such microfossils were uncovered from the Ruyang Formation in Shanxi, China and Roper Group, Northern Territories, Australia, dating back to 1.6-1.0 Ga ago. Some of these unicellular organic-walled fossils share characters with Ediacaran and Phanerozoic fossils, as well as extant green microalgae. Key characters among some Precambrian acritarchs are acetolysis-resistant vesicle with multi-layered walls; vesicle ornamentation by diverse processes that are produced during cyst formation; and excystment openings for the release of gametes or daughter-cells. Combination of these morphological elements, also present in extant phytoplankton, reflects the fossils’ protective function as reproductive cysts, indicating that complex life cycles and reproduction were well under way in Mesoproterozoic. Several case studies of microfossil morphology likely induced by intrinsic eukaryotic mechanisms are presented.Distinctive vesicle wall composed of the primary layer reinforced by polygonal platelets in Mesoproterozoic taxa Dictyosphaera and Shuiyouisphaeridium, as well as the sophisticated vesicle-wall patterning on the fossil sphaeromorphs Valeria and younger Cerebrosphaera would have required a certain degree of complexity for their formation, as observed in the present day analogues among eukaryotic protists. This suggests the activity of the key eukaryotic organelles and cellular mechanisms and signalling for the cyst formation. Considering that Golgi apparatus and the endoplasmatic reticulum are the organelles regulating eukaryotic secretory pathway and synthesis of biopolymers used in cell-wall construction, they would have been required for the complex morphology observed in these Precambrian taxa. Therefore, the presence of GA and ER in the eukaryotic cell is inferred at the minimum age of 1.6-1.4 Ga. Similarly, morphology of acritarchs of the Cambrian galeate plexus, namely openings with opercula, is likely induced by the activity of the LFA organelle (lid-forming apparatus) as in the extant dasycladalean alga Acetabularia.Additionally, several new morphotypes from the Ruyang Formation are presented. These unicellular fossils bear a velutinous outer membrane surrounding an internal sphere, which suggests a protective function of a reproductive or a resting cyst.Cyst-like morphology varies in disparity, but its key features are consistent through Mesoproterozoic, Neoproterozoic and early Palaeozoic.
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| 7. |
- Agić, Heda, 1989-, et al.
(författare)
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Organically-preserved multicellular eukaryote from the early Ediacaran Nyborg Formation, Arctic Norway
- 2019
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Ingår i: Scientific Reports. - : NATURE PUBLISHING GROUP. - 2045-2322. ; 9
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Tidskriftsartikel (refereegranskat)abstract
- Eukaryotic multicellularity originated in the Mesoproterozoic Era and evolved multiple times since, yet early multicellular fossils are scarce until the terminal Neoproterozoic and often restricted to cases of exceptional preservation. Here we describe unusual organically-preserved fossils from mudrocks, that provide support for the presence of organisms with differentiated cells (potentially an epithelial layer) in the late Neoproterozoic. Cyathinema digermulense gen. et sp. nov. from the Nyborg Formation, Vestertana Group, Digermulen Peninsula in Arctic Norway, is a new carbonaceous organ-taxon which consists of stacked tubes with cup-shaped ends. It represents parts of a larger organism (multicellular eukaryote or a colony), likely with greater preservation potential than its other elements. Arrangement of open-ended tubes invites comparison with cells of an epithelial layer present in a variety of eukaryotic clades. This tissue may have benefitted the organism in: avoiding overgrowth, limiting fouling, reproduction, or water filtration. C. digermulense shares characteristics with extant and fossil groups including red algae and their fossils, demosponge larvae and putative sponge fossils, colonial protists, and nematophytes. Regardless of its precise affinity, C. digermulense was a complex and likely benthic marine eukaryote exhibiting cellular differentiation, and a rare occurrence of early multicellularity outside of Konservat-Lagerstatten.
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| 8. |
- Agić, Heda, 1989-
(författare)
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Palaeobiology and diversification of Proterozoic-Cambrian photosynthetic eukaryotes
- 2015
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- One of the most important events in the history of life is the evolution of the complex, eukaryotic cell. The eukaryotes are complex organisms with membrane-bound intracellular structures, and they include a variety of both single-celled and multicellular organisms: plants, animals, fungi and various protists. The evolutionary origin of this group may be studied by direct evidence of past life: fossils. The oldest traces of eukaryotes have appeared by 2.4 billion years ago (Ga), and have additionally diversified in the period around 1.8 Ga. The Mesoproterozoic Era (1.6-1 Ga) is characterised by the first evidence of the appearance complex unicellular microfossils, as well as innovative morphologies, and the evolution of sexual reproduction and multicellularity. For a better understanding of the early eukaryotic evolution and diversification patterns, a part of this thesis has focused on the microfossil records from various time periods and geographic locations. Examination of microfossil morphology, cell wall microstructure and biochemical properties, reflect their intracellular complexity and function, and allow reconstructions of their life cycle, as well as observing the evolutionary pattern of change from Mesoproterozoic, to Cambrian-Ordovician transition. Several case studies included assemblages deriving from Mesoproterozoic, Neoproterozoic and early Paleozoic time intervals that show disparate morphotypes and innovative features indicative of algal clades. The Mesoproterozoic Ruyang Group in northern China has yielded a diverse microfossil assemblage that provides important clues about the diversification of different eukaryotic groups. Furthermore these microfossils contributed an additional evidence for the emergence of the crown group Eukarya by 1.7-1.4 Ga. In another part of this thesis, examination of wall microstructure and chemical properties via Raman spectroscopy has been used to assess the biological affinities of various Neoproterozoic problematic carbonaceous compression fossils. Studies on the early Phanerozoic (c. 545-485 Ma) assemblages from Estonia reconstructed patterns of the early radiations of phytoplankton and its evolutionary innovations. A continuing theme in this thesis has been using a combination of evidence of microfossils’ fine-scale morphology, ecology and chemical properties to determine their function in life, in addition to their systematic position.
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| 9. |
- Agić, Heda, 1989-, et al.
(författare)
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Raman spectra analysis and comparison of Neoproterozoic organic-walled mesofossils
- 2012
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Ingår i: The 2012 Fermor Meeting of the Geological Society. - London. ; , s. 86-
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Konferensbidrag (refereegranskat)abstract
- Skeletal material first appears in the rock record in the terminal Ediacaran, leaving most of the Earth’s history only to minute organic fossils. Aside from abundant acritarchs (unicellular organic-walled phytoplankton) present from at least ~2.5 Ga, other Proterozoic organic fossils of complex (eukaryotic) organisms include fungi and macroscopic algae of still debated taxonomy. Often preserved as flattened carbonaceous filaments in several morphologies: (1) rounded, (2) stick-like elongate and (3) branching, these Neoproterozoic fossils, including Chuaria, Morania, Beltanelloides, Vendotaenia, possess limited characters and differ little in gross morphology. As a result, other methods are needed to elucidate their biological affinities and, ultimately, phylogeny.Here we present the comparison of the Raman spectra analysis of different macroalgal genera from Yakutia, Siberia, as well as that of a putative polychaete Sabellidites from the East European Platform, dated to the early Cryogenian (840-700 Ma) and lowermost Cambrian respectively.Data from the vibrational modes of organic molecules from the wall of unmineralised organisms reveal their chemistry and partially wall ultrastructure, presumably an indication of their relationships. Polyaromatic chain hydrocarbons and n-aliphatic pyrolysates suggest algal affinity for some of the Neoproterozoic organic problematica, yet most of the Raman spectra results are still difficult to fully identify, partially owing to the thermal maturity of the host rocks. However, there are clear differences between various groups, differentiating between parts of a single plexus (cf previous studies of Chuaria-Tawuia suggesting them to be components of a multicellular plant) and elements from other taxa. Additionally, the distinct organic matter spectra of Chuaria and Sabellidites indicate that Raman spectroscopy could be a useful method in identifying different branches of the early eukaryotes.As they are usually shallow-water and dependant on sunlight, the record of sturdy photosynthetic macroorganisms in the Neoproterozoic strengthens the evidence that limits the extent of the harsh environmental conditions during the Cryogenian period, at least during the Kaigas and Sturtian glaciations.
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| 10. |
- Agić, Heda, 1989-, et al.
(författare)
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Raman spectroscopy and microstructural comparison of carbonaceous compression and body fossils from the Neoproterozoic of Siberian and Eastern European platforms
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Annan publikation (övrigt vetenskapligt/konstnärligt)abstract
- Macroscopic, organic-walled fossils preserved as carbonaceous compressions and body fossils are commonly occurring in the Neoproterozoic-Cambrian successions worldwide. Most of these fossils, including studied here Chuaria, Tawuia, and Beltemelliformis, have been accepted as algae, and Sabellidites as an early metazoan. They possess limited characters for biological identification and differ in gross morphology of spherodial vs. tubular millimetre-sized specimens. Consequently, other methods than morphologic observations are needed to elucidate their affinities and, ultimately, phylogeny. Here we present a comparison of the Raman spectrographic signatures and new scanning electron microscope (SEM) observations on different carbonaceous compression and body fossils from the Khajpakh Formation (Siberian Platform), and Nekrasovo Formation (East European Platform), referred to the Tonian-Cryogenian transitional interval (c. 840-700 Ma) and the lowermost Cambrian stage, respectively. Data from the Raman spectroscopy of the walls of non-mineralised organisms reveal their chemical properties, and, in additions to microstructural characters, may be used to resolve the fossils’ phylogenetic affinities. To test the basic recognition of organic matter in studied photosynthetic organisms vs. animals, we have examined algal compression fossils and organically-preserved body-fossil. Differences in the Raman spectroscopic signature between various taxa have been observed. Vibrational absorption bands similar to those characteristic of α-chitin signature have been detected in the organic wall of Sabellidites, consistent with its metazoan identity. Distinct organic matter spectra of the macroalgae Chuaria, Tawuia and Beltanelliformis, and the possible early annelid Sabellidites indicate that Raman spectroscopy could be a useful method in identifying different branches of the early eukaryotes. Additionally, the recognition of the earliest metazoans among un-diagnostic tubular fossils by biochemical signatures and wall ultrastructure, could provide the minimum age of their origins.
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