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A new conifer from the Upper Triassic of Southern Poland linking the advanced voltzialean type of ovuliferous scale with Brachyphyllum/Pagiophyllum -like leaves
Abstract
The Triassic is an exceptionally interesting period from the point of view of conifer evolution. The oldest representatives of some modern families appear in the fossil record in the Upper Triassic and all recent conifer families probably originated during this time. The stages whereby the primitive Voltziales transformed into the evolutionarily advanced families of modern conifers in the Triassic are yet to be fully documented. A newly discovered Polish Upper Triassic (Norian) locality in Patoka (SW Poland, Upper Silesia) with a unique fossil record offers us a rare opportunity to fill this gap. In this paper, we describe the very well preserved remains of a new conifer from this locality. We have been able to reconstruct the whole plant and propose a new taxon, Patokaea silesiaca gen. et sp. nov., on the basis of organic attachment and similar cuticular details on leaves, female cones, seed scale-bract complexes, ovules and developed seeds as well as male cones with pollen grains of the Enzonalasporites type in situ. This plant combines shoots bearing Brachyphyllum/Pagiophyllum-type leaves with a new type of seed scale-bract complex clearly derived from evolutionarily advanced Voltziales and male cones somewhat similar to the Cheirolepidiaceae (Classostrobus) type. Based on this distinctive and hitherto unknown feature combination, a new conifer family – Patokaeaceae – has been proposed. The female cones of this conifer were lax and borne singly at the end of leafy twigs. Ovulate scales were stalked and trilobate with two lateral oval lobes, each bearing one ovule, and one sterile reduced lobe between them, all in one plane. The bract was small and leaf-like. Male cones were simple, with microsporophylls helically arranged. Numerous pollen sacs were arranged around the microsporophyll stalk. Male cones were borne singly at the end of leafy twigs. This is also the first evidence of a relationship between Enzonalasporites pollen grains and previously unknown affinities, now found in situ in the male cones, ovule micropyle and inside the seed of this new conifer. This plant expands our view of the voltzialean conifer diversity lying at the root of modern conifer families.
... Remarks: All cuticles from Type E represent Brachyphyllum-like foliage. Brachyphyllum-foliage is a leaf morphogenus recognised from Triassic to Cretaceous and associated with different conifer groups: Cupressaceae, Podocarpaceae, Araucariaceae, Cheirolepidiaceae (Meyen, 1987;Stockey, 1994) and Patokaeceae (Pacyna et al., 2017). Type E cuticles show gross characteristics of Cheirolepidiaceae family (i.e., random orientation of stomata, 4 to 6 subsidiary cells and distinctive papillae on them; Watson, 1988;Ash, 1999;Bomfleur et al., 2011). ...
... i.e., Pagiophyllum-like (Pacyna et al., 2017) within one fossil plant. Investigated cuticles are not classified as Brachyphyllum sp., but as a separate cuticulae dispersae morphotype. ...
... However, the ex situ character of the sediments presents an obstacle to a comprehensive palaeoecological reconstruction. Nevertheless, the studied fossils give a general picture of the Norian plant assemblage, consistent with other observations on Norian floras from Northern Hemisphere (Dobruskina, 1994;Pacyna et al., 2017;Kustatscher et al., 2018;Pacyna, 2019). Generally, the dry Norian, favoured xerophytic plants, but dry-humid climate fluctuations (Szulc et al., 2006) facilitated the growth of other plant ecogroups. ...
... Taxa from Kühwiesenkopf/Monte Prà della Vacca discussed in this article are indicated with an asterisk. 1851-1852Frentzen 1931;Grauvogel-Stamm and Grauvogel 1975;Grauvogel-Stamm 1978;Miller 1982;Rothwell et al. 2000;Gall and Grauvogel-Stamm 2005;Díez et al. 2010;Escapa et al. 2010;Herrera et al. 2015;Pacyna et al. 2017 Olenekian-early Anisian Ortiseia leonardii Florin, 1964Florin 1964Clement-Westerhof 1984, 1987, 1988Kustatscher et al. 2012 Wuchiapingian Pachylepis: Kräusel, 1952Linck 1950Kräusel 1952 Pachylepis quinquies (Linck) Kräusel, 1952Linck 1950Kräusel 1952;Grauvogel-Stamm 1978;Kelber and Hansch 1995;Axsmith and Taylor 1997 Late Ladinian-Carnian 2002Kräusel 1939Mägdefrau 1956Mägdefrau , 1963Arndt 2002;Pacyna et al. 2017 Carnian Coburg, Hassfurt, Germany ...
... Taxa from Kühwiesenkopf/Monte Prà della Vacca discussed in this article are indicated with an asterisk. 1851-1852Frentzen 1931;Grauvogel-Stamm and Grauvogel 1975;Grauvogel-Stamm 1978;Miller 1982;Rothwell et al. 2000;Gall and Grauvogel-Stamm 2005;Díez et al. 2010;Escapa et al. 2010;Herrera et al. 2015;Pacyna et al. 2017 Olenekian-early Anisian Ortiseia leonardii Florin, 1964Florin 1964Clement-Westerhof 1984, 1987, 1988Kustatscher et al. 2012 Wuchiapingian Pachylepis: Kräusel, 1952Linck 1950Kräusel 1952 Pachylepis quinquies (Linck) Kräusel, 1952Linck 1950Kräusel 1952;Grauvogel-Stamm 1978;Kelber and Hansch 1995;Axsmith and Taylor 1997 Late Ladinian-Carnian 2002Kräusel 1939Mägdefrau 1956Mägdefrau , 1963Arndt 2002;Pacyna et al. 2017 Carnian Coburg, Hassfurt, Germany ...
... One of the characteristics of the Triassic voltzialean pollen cones (e.g., Aethophyllum, Darneya, Hercynostrobus, Ruehleostachys, Sertostrobus, Willsiostrobus;e.g., Roselt 1956;Schaarschmidt and Maubeuge 1969;Grauvogel-Stamm 1978;Grauvogel-Stamm and Schaarschmidt 1979;Arndt 2002;table 4; fig. 15) compared with other younger conifer forms (e.g., Classostrobus, Hirmeriella, Kobalostrobus; e.g., Serlin et al. 1981;Krassilov 1982b;Clement-Westerhof and Van Konijnenburg-van Cittert 1991;Barbacka et al. 2007;Rothwell et al. 2007;Hieger et al. 2015;Pacyna et al. 2017) is their larger size. The size of polliniferous cones was gradually reduced from the most ancient to the younger forms, with some exceptions in the Upper Triassic and Lower Cretaceous (e.g., Patokaea, Tomaxellia; Archangelsky and Gamerro 1967;Archangelsky 1968;Pacyna et al. 2017). ...
... All previous ultrastructural studies on fossil (cheirolepid) conifer pollen grains have used transmitted electron microscopy (TEM) (e.g. Pacyna et al., 2017;Taylor and Alvin, 1984;Zavialova et al., 2010a;Zavialova and Roghi, 2005;Zavialova et al., 2010b). FIBSEM is an effective and, for some tasks, superior alternative to TEM (Villanueva-Amadoz et al., 2012) as it produces essentially 3D images which can be viewed sequentiallylike a movie (Supplementary Video File 1)or from different angles, whereas TEM produces isolated, static 2D sections. ...
... Diverse views on the morphology of Enzonalasporites group genera were published following Leschik (1955). Enzonalasporites and Patinasporites have variously been considered saccate alete spores (Boulouard, 1963;Klaus, 1960;Leschik, 1955;Potonié, 1966;Scheuring, 1970), monosaccate trilete spores (Biard, 1963), velate pollen without recognizable germinalia (Mädler, 1964), velate pollen with a sulcus (Schulz, 1967) and asaccate pollen (Balme, 1995;Pacyna et al., 2017). The curved walls of rugulae transversely crossing the corpus in Vallasporites were interpreted as a bulging trilete mark (Scheuring, 1970), not a trilete mark (Potonié, 1958), or not commented on (Leschik, 1955). ...
... Unlike the images from Pacyna et al. (2017) which indicate Enzonalasporites pollen grains are asaccate (their fig. XII, [3][4][5], our FIBSEM images show there is inflation of the exine laterally in some specimens (Plate IV,5;Plate V,4). ...
The ‘Enzonalasporites group’ comprises Enzonalasporites, Patinasporites, Vallasporites, Pseudenzonalasporites, Tulesporites, Daughertyspora and Zonalasporites pro parte, and their described species. There is little consensus on the taxonomy of this widely distributed group of mainly Late Triassic pollen despite their importance in biostratigraphy, paleoecology and conifer evolution. Last revised over forty years ago, their taxonomy is plagued by inadequate diagnoses and emendations, inconsistent or inaccurate terminology, and intergrading morphological attributes. In this paper we present new data and interpretations on the morphology, ornamentation and ultrastructure of key species, and use them together with analysis of published images and data to underpin a major review of the group's taxonomy and position in conifer evolution. Confocal laser scanning microscopy (CLSM) shows the raised ridges of rugulae on the proximal face of some pollen grains are impression marks, not trilete marks. Focused ion beam scanning electron microscope (FIBSEM) sectioning of some E. vigens and Patinasporites densus pollen grains shows that overlapping and intertwined rugulae create alveoli in a slightly inflated lateral exinal envelope, or “saccoid”. Dimensions of the corpus, saccoid and rugulae were measured using both SEM and transmitted light microscopy (LM) images. The challenge of intergrading attributes was managed by selecting morphologically stable criteria that can be used with routine LM, an approach which enables most specimens to be assigned taxonomically with confidence, while acknowledging that some may remain problematic. The hierarchy of criteria used in revising the taxonomy were: (1) corpus diameter/total width ratio, (2) dominant sculptural element, (3) presence or absence of impression marks, and (4) saccoid width and characteristics. The first two criteria help discriminate between genera, and the second two between species. After applying these criteria, emending diagnoses and synonymizing or transferring several species, three genera and seven species remain: E. vigens, E. ignacii, E. antonii n.c., Patinasporites densus, Pseudenzonalasporites cinctus n.c., Pseudenzonalasporites leschikii n.c., and Pseudenzonalasporites summus. The Enzonalasporites group reached maximum diversity, abundance and distribution during the Carnian, a stage characterized by several humid episodes, but the details of their paleobiogeographic distribution are complex. E. vigens and E. ignacii seem to have preferred xerophytic or halophytic conditions with occasional wetter episodes (e.g. monsoons), while Patinasporites densus seems to have preferred more hygrophytic conditions. The common thread between these species is their requirement for moisture during germination which required a pollination drop. Paleoenvironmental changes during the latest Triassic may have favored conifers which germinated using siphonogamy.
... Patokaea Pacyna, Barbacka et Zdebska, a representative of a new voltzialean family, was described from the Norian of Silesia (Pacyna et al., 2017). The in situ pollen grains are medium in size (average size is 32-35 μm), circular in polar view to oval in equatorial or oblique view, with a rugulate surface and without sacci; neither proximal scar nor distal aperture was detected. ...
... Carrière and to Cerebropollenites; the endexine is very thin, fine-grained, more electron-dense than the ectexine. Similar to us, Pacyna et al. (2017) also faced difficulties in finding a corresponding dispersed taxon. Most closely comparable appeared Enzonalasporites Leschik emend. ...
... Pollen grains that are conventionally incorporated in these taxa are more or less rounded and medium in size; there is no general agreement in the interpretation of their exine architecture: monosaccate, monosaccoid, velate and asaccate states were named (see Pacyna et al., 2017 for references). Vallasporites differs by a bulging proximal trilete scar that reaches the equator; Pseudoenzonalasporites, by a different exine pattern and distinctly dark inner body; and Patinasporites, by a wide ectexine that can be described as an equatorial zone or saccus (Pacyna et al., 2017). ...
The amount of information about the diversity of dispersed pollen grains and spores exceeds that about their in situ counterparts by several orders of magnitude. Strangely, there are several finds of in situ pollen types that are thus far unknown from dispersed palynomorph assemblages. Here we provide a brief discussion of this peculiar phenomenon based mainly on two examples from our recent studies, including a late Permian lyginopterid seed fern from Jordan and a Middle Jurassic conifer from Siberia. Several reasons could account for such “in situ only” pollen types, including the scarcity of the parent plants, low pollen productivity, entomophily, or immaturity of in situ pollen, although the last variant seems overestimated. Such pollen grains, when found dispersed, might also be considered as untypical, featureless or inadequately preserved specimens and, as a consequence, be reported without illustrations as undetermined specimens or even ignored altogether; this way, they receive little attention from the authors, remain unregistered, and cannot be analyzed by subsequent researchers. We stress the importance of publication of as complete as possible information about all components of palynological assemblages, including their photos, even if they do not belong to stratigraphically important taxa. We do not attribute the disparity to taphonomic differences between megafossil and microfossil deposition, because the ‘in situ only’ pollen types are absent not only in the same beds where the pollen organs were found, but are unknown from any other deposits.
... Cheirolepidiaceae was dominant along the banks of rivers and lakes. During this period, the remains of the voltzialean Patokaea silesica with polliniferous cones containing pollen grains of the genus Enzonalasporites were found in the fluvial-lacustrine deposits of Patoka (Upper Silesia) (Pacyna et al., 2017). ...
... They were found inside the micropylar canal of the ovule (Figure 2A), at the micropylar end of the seed, and in the pollen sacs of polliniferous cones ( Figure 2B,C). This is not only the first evidence of a relationship between Enzonalasporites pollen and the parent plant (Pacyna et al., 2017) but also a rare and valuable record of pollen grains found inside the micropylar canal of the ovule, fossilized at the precise moment of pollination. ...
Palynological studies of the Mesozoic era in Poland began in the 1950s. These investigations developed in many directions, including stratigraphy, systematics of spores and pollen grains and their botanical affinities, as well as paleoecological and paleoclimatic interpretations. In this study, we provide an overview of the most important achievements in Mesozoic terrestrial palynology in Poland.
... Remarks: Enzonalasporites Leschik, Patinasporites Leschik, Pseudoenzonalasporites Scheuring, and Vallasporites Leschik are comparable to the in situ pollen of Patokaea Pacyna, Barbacka et Zdebska (Pacyna et al., 2017). Some authors describe this type of pollen as monosaccate or monosaccoid pollen (Van der Eem, 1983). ...
... Some authors describe this type of pollen as monosaccate or monosaccoid pollen (Van der Eem, 1983). As the pollen grains that found in a Patokaea silesiaca Pacyna, Barbacka et Zdebska, show no alveolar structure under TEM, they are probably asaccate (Pacyna et al., 2017). However, detailed TEM investigations are also needed for the dispersed pollen grains related to this type of pollen. ...
The ecogroup classification based on the growth-form of plants (Eco-Plant model) is widely used for extant, Cenozoic, Mesozoic, and Paleozoic paleoenvironmental reconstructions. However, for most Mesozoic dispersed sporomorphs, the application of the Eco-Plant model is limited because either their assignment to a specific ecogroup remains uncertain or the botanical affinities to plant taxa are unclear. By comparing the unique outline and structure/sculpture of the wall of dispersed sporomorph to the sporomorph wall of modern plants and fossil plants, 861 dispersed Mesozoic sporomorph genera of Bryophytes, Pteridophytes, and Gymnosperms are reviewed. Finally, 474 of them can be linked to their closest parent plants and Eco-Plant model at family or order level. Based on the demands of the parent plants to different humidity conditions, the Eco-Plant model separates between hydrophytes, hygrophytes, mesophytes, xerophytes, and euryphytes. Additionally, due to different temperature demands a separation in megathermic, mesothermic, microthermic, and eurythermic plants is possible. In the Mesozoic, both spore-producing and pollen-producing plants are adapted to different kinds of humidity. The concept to use the spore/pollen ratio to reflect the hygrophytes/xerophytes ratio is therefore questionable. The presented ecogroups for dispersed Mesozoic sporomorphs now allow identifying at least relative plant, paleoenvironmental and paleoclimate changes in Mesozoic sedimentary records.
... Unlike S. gracile, Triassic pollen cones of Patokaea silesiaca Pacyna, Barbacka et Zdebska (Pacyna et al. 2017) are small (3.8-6.0 mm long and 1.8-4.0 mm wide) and characterized by triangular to wedge-shaped (in a compressed state) pollen sacs that were probably sessile and arranged in a whorl around the central stalk of the microsporophyll (table A1). ...
... Our pollen grains are similar to this group in their more or less rounded outlines, occasional presence of a small rudimentary proximal scar, and fine but distinct sculpturing. Thus, a recent study by Pacyna et al. (2017) revealed circular pollen grains with distinct sculpturing in an Upper Triassic plant of a presumably voltzialean affinity, P. silesiaca. In search of a dispersed analogue, the authors chose from among Enzonalasporites Leschik emend. ...
Premise of research. Jurassic localities of fossil flora in the Irkutsk Region (East Siberia) are rich in reproductive remains of gymnosperms with in situ pollen. Our aim was to understand the morphology of pollen grains from conifer cones of Schidolepium gracile in the context of the botanical affinity of their parent plant.
Methodology. Pollen grains were studied in transmitted light as well as with SEM and TEM.
Pivotal results. The pollen grains demonstrate an unusual combination of morphological and ultrastructural traits. In pollen masses, they appear circular, asaccate, and flattened in a polar position. Detached monads show a polar and equatorial position equally often; the outlines are rounded, oval, and irregular. An equatorial-distal saccus was revealed. A small trilete scar is occasionally present. The surface is fossulate. The ectexine is formed by structural elements that fused with each other by their lateral surfaces partially or completely. The element is a solid elongated cylinder with a rounded external end and narrowing internal end/ends. The elements are arranged along their length, perpendicular to the pollen surface. The endexine is more electron dense than the ectexine, and it is prominent and appears homogeneous. We revealed variations in pollen morphology that we consider preservational, although the existence of two species of Schidolepium is not excluded. One of the cones was contaminated by pollen grains of the Cycadopites type; electron microscopical data proved their ginkgoalean affinity.
Conclusions. The exine ultrastructure excludes an araucariaceous affinity, in spite of a relatively close general morphology. Certain similarities to Cerebropollenites were found in the exine ultrastructure, but it possesses a distal aperture and proximal saccus-like extensions. Although the pollen cones have something in common with the Taxodiaceae and Voltziales, the palynological data do not support this relationship. The plant apparently represents an early member of an evolutionary line within conifers.
... By contrast, the pollen grains reported here are flattened in various positions. Enzonalasporites is sometimes interpreted as monosaccate (Van der Eem 1983) and sometimes as asaccate (Pacyna et al. 2017). Enzonalasporites is medium-sized and shows a surface pattern similar to the present material, but lacks an alveolate ectexine (Pacyna et al. 2017). ...
... Enzonalasporites is sometimes interpreted as monosaccate (Van der Eem 1983) and sometimes as asaccate (Pacyna et al. 2017). Enzonalasporites is medium-sized and shows a surface pattern similar to the present material, but lacks an alveolate ectexine (Pacyna et al. 2017). Kuglerina, Doubingerispora, Cordaitina, Parasaccites, and Perisaccus all differ from the present form in showing clearly developed sacci. ...
Here we report a pollen organ from the upper Permian of the Dead Sea region with in situ pollen showing a wall structure typical for lyginopterid seed ferns. The single specimen resembles the dispersed seed-fern pollen organs Dictyothalamus and Melissiotheca in being composed of numerous densely positioned, radiating pollen sacs that together form a characteristic honeycomb-like surface pattern. The small, rounded to oval, non-saccate cryptaperturate in situ pollen grains have a verrucate sculpture and show an alveolate wall ultrastructure with alveoli arranged in several tiers. The lyginopterids, a group of seed ferns, first appeared in the latest Devonian and were very widespread across Euramerica during the Mississippian. The group shows a clear demise in the middle and late Pennsylvanian. The youngest representatives have been reported from late Permian peat-swamps in Cathaysia. The morphology of the pollen organ, the ultrastructure of the pollen, and associated foliage present further evidence that lyginopterid seed ferns persisted until the late Permian and demonstrate that they were not restricted to peat-forming swamp environments, but that they were able to survive in drier environments.
... We suggest that this increase in metabolic potential fostered the rapid rates of evolutionary change found among crown mammals during this period, and so may have facilitated the Early to Middle Jurassic adaptive radiation of crown mammals. A new conifer family, Patokaeaceae, comprising one genus and species, Patokaea silesiaca, has been recently described from the Upper Triassic of southern Poland (Pacyna et al., 2017). The plant remains are preserved as coalified compressions with well-preserved cuticles. ...
... Also, round holes are sometimes visible on the leaf cuticle, although are absent from mined specimens. Several shoot apexes differ from the typical condition described for this species by Pacyna et al. (2017). Altered, atypical shoot apexes are shortened, swollen, and their leaves are very densely arranged. ...
... We suggest that this increase in metabolic potential fostered the rapid rates of evolutionary change found among crown mammals during this period, and so may have facilitated the Early to Middle Jurassic adaptive radiation of crown mammals. A new conifer family, Patokaeaceae, comprising one genus and species, Patokaea silesiaca, has been recently described from the Upper Triassic of southern Poland (Pacyna et al., 2017). The plant remains are preserved as coalified compressions with well-preserved cuticles. ...
... Also, round holes are sometimes visible on the leaf cuticle, although are absent from mined specimens. Several shoot apexes differ from the typical condition described for this species by Pacyna et al. (2017). Altered, atypical shoot apexes are shortened, swollen, and their leaves are very densely arranged. ...
... Protocupressinoxlon carrizalense, the taphoflora includes voltzian leaves of Heidiphyllum elongatum and pollen grains allied to conifers, such as Enzonalasporites (Césari et al., 2021), a genus related to the family Patokaeaceae, which combines voltzian and cheirolepidiacean characters (Pacyna et al., 2017). ...
... e shape and size of the pollen cones resemble those of Classostrobus, the male cone of Jurassic-Cretaceous Cheirolepidiaceae, but the arrangement of the pollen sacs on the microsporophyll corresponds more closely with that of Voltziales. It seems that this family did not continue to evolve further within conifers but turned out to be important in evolutionary considerations (Atkinson et al., 2018;Leslie et al., 2018) and in research on pollen preserved in situ (for references see Pacyna et al., 2017;Zhang et al., 2021). is is the first and so far the only detailed documented fossil coniferous plant in Polish paleobotany. ...
Research on Polish Mesozoic macroflora has remarkably changed over the last 100 years. Compared with traditional nineteenth-century taxonomical and evolutionary studies, modern research also investigates of fossil plant diversity, plant interactions with the environment and animals, and deep changes in vegetation influenced by environmental and climate change. In this review, we present the most important turning points in Polish Mesozoic macroflora research and the development of cognitive techniques by discussing selected studies carried out by Polish palaeobotanists.
... While FIBSEM has been used in materials science and the semiconductor industry for several decades, it has only been used infrequently in palynology (e.g., (House and Balkwill, 2013, 2016Lupia et al., 2015;Schiffbauer and Xiao, 2009;Villanueva-Amadoz et al., 2012), and only once on conifers (Villanueva-Amadoz et al., 2014). Ultrastructural studies of cheirolepid pollen have used transmitted electron microscopy (TEM) to date (Médus, 1977;Taylor and Alvin, 1984;Litwin and Skog, 1991;Pacyna et al., 2017;Zavialova et al., 2010a;Zavialova and Roghi, 2005;Zavialova et al., 2010b); of these, only Litwin and Skog (1991) examined Camerosporites pollen grains. FIBSEM is an effective and, for some tasks, superior alternative to TEM (Villanueva-Amadoz et al., 2012) as it produces essentially 3D images which can be viewed sequentiallylike a movieor from different angles, whereas TEM produces isolated, static 2D sections. ...
This paper presents new morphological and ultrastructural data on Camerosporites pollen obtained from a range of microscopy techniques, and uses these (and other) data to emend the taxonomy of the genus, and review aspects of the paleobiology of its parent plants and their significance in conifer evolution. Camerosporites, a circumpolloid genus probably produced by cheirolepids, was most abundant and widely distributed in low to middle latitudes during the Carnian before declining and becoming extinct in the latest Triassic or perhaps Early Jurassic. The Cheirolepidiaceae, a large and diverse family of early conifers, are characterized by their unique combination of pollen attributes including a rimula, polar tenuitates, exinal threads, verrucae, tetrads/dyads, and exinal and sculptural variability. Confocal laser scanning microscopy (CLSM) and scanning electron microscopy (SEM) images show most C. secatus pollen grains have a clear proximal rimula and some a probable distal rimula. No convincing specimens with two rimulae were seen. Images from focused ion beam scanning electron microscopy (FIBSEM) show that the verrucae are essentially hollow chambers with thickened outer walls and floored by an irregular reticulate-like network of fine, elongated to granular exinal elements. These images suggest C. secatus pollen grains have relatively “primitive” ultrastructure for a circumpolloid – homogenous nexine, (some) granulae in the infratectum, and an indistinct or absent tectum. Camerosporites was able to thrive in the periodically humid Carnian, clearly demonstrating the vegetative organs of its parent plants were able to adapt to a wide range of paleoenvironments. Nevertheless, Camerosporites may have been out-competed for increasingly scarce water resources in the drying Norian-Rhaetian by other producers of circumpolloid pollen and early conifers with distinct pollination strategies possibly related to siphonogamy. We synonymized C. pseudoverrucatus and C. spissus with C. secatus as they are hard to differentiate consistently given their variability and intergrading attributes. We retained C. verrucosus and, tentatively, C. reductiverrucatus and C. hengyangensis. C. reductiverrucatus can plausibly be interpreted as reworked and degraded C. secatus rather than a different species.
... However, also the families Podocarpaceae (Samaropollenites), Araucariaceae (Araucariacites), Pinaceae (Abietinaepollenites) are represented in the palynological record. The monosaccate pollen grains are assigned to the Voltziales but without any detailed botanical affinity since the so far only in situ record is that of the putative voltzialean Patokaea silesiaca (Pacyna et al., 2017). Among the spores dominate those of the lycophytes (4/10 genera) and the ferns (4/10 genera). ...
The Carnian Pluvial Episode (CPE) has been recognized as a time of plant radiations and originations, likely related to observed swift changes from xerophytic to more hygrophytic floras. This suggests that the increasing humidity causally resulting from LIP volcanism was the trigger for these changes in the terrestrial realm. Understanding the cause and effects of the CPE on the plant realm, requires study of well-preserved floras that are precisely aligned with the CPE. We therefore focus on the best age-constrained section within the CPE for the terrestrial to marginal marine environment to understand the floristic composition at the early CPE. This is found in the Dolomites, Italy, and is remarkable for the preservation of the oldest fossiliferous amber found in the rock record. An integrated study of palynomorphs and macro-remains related to the conifer families of the fossil resin bearing level brings together the floral components from this section. This observed mixture of different taxa of extinct and modern conifer families underlines firmly the effects of the LIP-induced CPE on the evolution and radiation of conifers.
... Recent discoveries of Upper Triassic outcrops in Silesia (southwestern Poland) have yielded not only rich vertebrate material, but also new plant fossils (Dzik and Sulej, 2007;Pacyna, 2014). Plant macrofossils rarely are found in the Upper Triassic of Poland (Reymanówna, 1986;Pacyna, 2014;Pacyna et al., 2017;Kustatscher et al., 2018) and hence are poorly known; such discoveries open new opportunities for study of them. Krasiejów was the first Polish Triassic site with the bones of large land vertebrates described and it is still one of the most important Lucas, 2018). ...
... Later conifers appear to have altered the development of the ovuliferous complexes, producing more reduced appendages on their fertile shoots. The socalled "voltzialean" or "transition" conifers of the Permian and Triassic (see Clement-Westerhof 1987;Looy 2007;Taylor et al. 2009;Pacyna et al. 2017) produced ovuliferous scales with lobes that are typically interpreted as the remains of leaves (Fig. 4). Extant clades emerged in the Late Triassic (Taylor et al. 2009;Leslie et al. 2012), and their ovuliferous scales typically lack any trace of appendages ( Fig. 4; although this reduction series may be complex and lineage-specific in reality; see Dörken and Rudall 2018). ...
Biologists often study morphological evolution through form and function relationships. But biological structures can perform multiple functional roles, complicating efforts to understand the evolutionary significance of any one relationship. Plant reproductive organs perform multiple roles in a sequence, however, which provides a unique opportunity to understand how structures evolve to meet multiple functional demands. Using conifers as a study group, we discuss how a shared developmental trajectory links the performance of sequential functional roles. Variation in development among lineages can underlie morphological diversity; pollination-stage seed cones in Pinaceae conifers function similarly but show diverse forms reflecting differences in developmental rate. As cones develop further, the morphologies that they use to perform later functional roles is influenced by the specific developmental patterns used to meet earlier demands, which may ultimately limit morphological diversity. However, we also show how selective pressures relating to the final functional stage (seed dispersal) may influence cone anatomy and morphology over all previous stages, highlighting their complex linkages among form, function, and development. We end by discussing the potential relationships between functional ontogeny and morphological disparity in plant reproductive structures more broadly, suggesting that the complex functional roles associated with seed plant reproduction probably underlies the high disparity in this group.
... Recent discoveries of Upper Triassic outcrops in Silesia (southwestern Poland) have yielded not only rich vertebrate material, but also new plant fossils (Dzik and Sulej, 2007;Pacyna, 2014). Plant macrofossils rarely are found in the Upper Triassic of Poland (Reymanówna, 1986;Pacyna, 2014;Pacyna et al., 2017;Kustatscher et al., 2018) and hence are poorly known; such discoveries open new opportunities for study of them. Krasiejów was the first Polish Triassic site with the bones of large land vertebrates described and it is still one of the most important ( Sues and Fraser, 2010;Lucas, 2018). ...
The Early Jurassic flora of the Mecsek Mountains is diverse, with numerous representatives of ferns, seed ferns, cycadophytes, ginkgophytes, and conifers. Its (para-)autochthonous deposition, good preservation and low collection bias has permitted researchers to save delicate, small plant remains that would generally be missing from the fossil record. These plant fossils are characterized by having a “filmy” (probably unilayered) structure, thin stalks with flat leaf-like branches and ultimate irregular segments with streamlined epidermal cells having thick cell walls. Due to the absence of sporangia, the plant remains cannot be confidently assigned to any higher plant group, although they show some similarities to thalloid liverworts with raised vegetative bodies and also have some resemblance to the fern family Hymenophyllaceae. The new genus and species Leonophyllum tenellum Barbacka et Kustatscher is erected.
... The abundance of recent diversification within otherwise widely divergent conifer lineages also limits their utility for reconstructing specific patterns of trait evolution, because large morphological branch lengths separate the surviving lineages. Paleobotany is therefore essential to understanding conifer phenotypic evolution, because it offers the potential to break up long morphological branches by identifying and characterizing extinct taxa with unusual traits (e.g., Herrera et al., 2015;Pacyna et al., 2017) and by testing the phylogenetic placement of such taxa (Escapa et al., 2012). FIGURE 5. Potential pathways of evolutionary change in conifer seed cones given the phylogenetic relationships suggested by molecular data. ...
Premise of the Study
Conifers are an important living seed plant lineage with an extensive fossil record spanning more than 300 million years. The group therefore provides an excellent opportunity to explore congruence and conflict between dated molecular phylogenies and the fossil record.
Methods
We surveyed the current state of knowledge in conifer phylogenetics to present a new time‐calibrated molecular tree that samples ~90% of extant species diversity. We compared phylogenetic relationships and estimated divergence ages in this new phylogeny with the paleobotanical record, focusing on clades that are species‐rich and well known from fossils.
Key Results
Molecular topologies and estimated divergence ages largely agree with the fossil record in Cupressaceae, conflict with it in Araucariaceae, and are ambiguous in Pinaceae and Podocarpaceae. Molecular phylogenies provide insights into some fundamental questions in conifer evolution, such as the origin of their seed cones, but using them to reconstruct the evolutionary history of specific traits can be challenging.
Conclusions
Molecular phylogenies are useful for answering deep questions in conifer evolution if they depend on understanding relationships among extant lineages. Because of extinction, however, molecular datasets poorly sample diversity from periods much earlier than the Late Cretaceous. This fundamentally limits their utility for understanding deep patterns of character evolution and resolving the overall pattern of conifer phylogeny.
... Furthermore, it is worth noting that little is known about the anatomy of extant and extinct araucariaceous pollen cones, thus making it difficult to compare Chimaerostrobus to Araucariaceae in any detail. A diversity of pollen sac arrangements has been recorded within extinct conifer groups (Grauvogel-Stamm, 1969, 1978Schaarschmidt and Maubeuge, 1969;Serlin et al., 1980;Hermsen et al., 2007;Hernandez-Castillo et al., 2009a, 2009bPacyna et al., 2017). Pollen sacs of some of these conifers such as Table 1). ...
Paleontological data indicates that the Late Triassic–Early Jurassic was a critical time interval for the phylogenetic and morphological diversification of conifers, especially modern families. In this study a new genus and species, Chimaerostrobus minutus, is characterized based on an anatomically preserved pollen cone from the Lower Jurassic of Antarctica. The cone was prepared using the cellulose acetate peel technique and studied using light and scanning electron microscopy. The pollen cone of Chimaerostrobus is subspheroidal with tightly imbricate microsporophylls. Microsporophylls contain a single vascular bundle and abundant transfusion tissue. There are 22 pollen sacs per microsporophyll that are both abaxially and adaxially attached to the stalk and distal laminar head. In situ pollen grains are oblate, 15–20 μm in equatorial diameter, and asaccate. Chimaerostrobus has a combination of characters indicative of Araucariaceae and extinct conifers such as voltzialeans and Kobalostrobus; however, due to its unique character mosaic it cannot be assigned to an established group. This new species sheds light on the complexity of pollen cone evolution and the diversity of conifers that were evolving during the Late Triassic and Early Jurassic.
The Carrizal Formation, exposed in the Marayes-El Carrizal Basin (western Argentina), has been the focus of palaeobotanical studies since the late 1800s. The recent finding of well-preserved palynological assemblages provides the first detailed studies about its palynofloras. In this paper, the 63 taxa identified in the unit are illustrated and discussed, as well as their stratigraphic distribution in equivalent palynological assemblages of Argentina. Some spore species are revised: Uvaesporites hammenii (Herbst) Césari, comb. nov., Retusotriletes wielandii (Jain) Césari, comb. nov. and Lundbladispora stellae (Herbst) Césari, comb. nov. The recognition of Cadargasporites baculatus de Jersey & Paten emend. Reiser & Williams, Craterisporites rotundus de Jersey, Enzonalasporites vigens Leschik, Leptolepidites argenteaeformis (Bolkhovitina) Morbey, Protodiploxypinus americus Dunay & Fisher and Rugulatisporites permixtus Playford, among others, appears to be useful for local and intercontinental correlations. A Carnian age is proposed for the palynofloras.
Three spore-pollen assemblages, representing Lundbladispora obsoleta-Protohaploxypinus pantii and Densoisporites nejburgii Zones, were distinguished within the Lower and Middle Buntsandstein deposits in NW part of the Holy Cross Mts. Palynofacies analysis provided for more detail data about the changes of depositional environment during the Early Triassic in the studied area. Palaeoenvironmental model used here indicates two climatic cycles in the Lower and Middle Buntsandstein. -Author
The current Triassic chronostratigraphic scale is a hierarchy of three series (Lower, Middle, and Upper) divided into seven stages (Lower = Induan and Olenekian; Middle = Anisian and Ladinian; and Upper = Carnian, Norian, and Rhaetian), further divided into 15 substages (Induan = upper Griesbachian and Dienerian; Olenekian = Smithian and Spathian; Anisian = Aegean, Bithynian, Pelsonian, and Illyrian; Ladinian = Fassanian and Longobardian; Carnian = Julian and Tuvalian; and Norian = Lacian, Alaunian, and Sevatian). Ammonoid and conodont biostratigraphies provide the primary basis for the chronostratigraphy. We argue here for reliance on ammonoid-based biostratigraphy for Triassic chronostratigraphic definitions. We advocate a four-stage Lower Triassic and the elevation of the very long Carnian and Norian stages to series. A sparse but growing database of precise radioisotopic ages supports the following calibrations: base of Triassic ~252 Ma, base Olenekian ~251 Ma, base Anisian ~247 Ma, base Ladinian ~242 Ma, base Norian ~221 Ma, and base Jurassic ~201 Ma. Triassic magnetostratigraphy is a series of multichrons at best, and needs vast improvement to make a serious contribution to the Triassic timescale.
Two conifers are briefly described and illustrated from a rich plant-bearing layer of late Carnian age from the middle New Oxford Fm., Gettysburg Basin, USA. Two types of leaf-bearing branch systems are assigned to Pagiophyllum diffusum (Emmons) nov. comb. and P. simpsonii Ash. Seed cones that I assign to Glyptolepis nov. sp.1 and cf. G. nov. sp. 2 were also found. Six cones of G. nov. sp. 1, bearing large ovoid winged seeds, were found attached to a branch bearing leaves of P. diffusum. Both types of leaves and cones possess the same hairy type of cuticle. A small, elongate pollen cone, also with a hairy cuticle, was recovered from the plant layer. This cone contains Patinasporites densus Leschik pollen. P. densus and a morphologically similar pollen type, Vallasporites ignacii Leschik, dominate the palynoflorule from the plant layer.
The stratigraphy of the Upper Silesian Keuper, a continental mudstone-dominated is poorly known, although the already renowned, newly-discovered vertebrate localities highlight a growing demand for a more precise intra-regional correlation and an appropriate stratigraphic context. A major lithostratigraphic unit, preliminarily proposed for the middle Keuper (i.e., above the Schilfsandstein; Stuttgart Formation in “Stratigraphische Tabelle von Deutschland”, 2002) by Szulc and Racki (2015; Przegląd Geologiczny, 63: 103–113), is described in detail. The re-defined Grabowa Variegated Mudstone-Carbonate Formation, based on previously inaccurately presented information, includes the Upper Gypsum Beds and Steinmergelkeuper in traditional scheme from Germany (= Weser and Arnstadt Formations). Three members are formally defined: Ozimek (mudstone-evaporite), Patoka (marly mudstone-sandstone) and Woźniki (limestone). Two significant bone-bearing horizons (Krasiejów and Lisowice) are placed within the Patoka Mbr. The formation thickness in a composite regional reference section of the Upper Silesian Keuper, based on the new Woźniki K1 and Patoka 1 well profiles, is approximately 230 m. The Grabowa Fm generally correlates with the Norian stage, with the base located in an undefined upper Carnian, and it is topped by a major erosive disconformity and sedimentary sequence boundary near the Norian-Rhaetian boundary. However, hiatuses in the Silesian middle Keuper succession are located and paired with a cannibalistic type of sand-mud flat deposition, largely controlled by Early Cimmerian tectonic block movements around the Kraków-Lubliniec shear zone.
The thematic issue, encompassing 8 articles, written by authors from nine Polish institutions, USA and Germany, is reviewed. This issue presents results of the project “The evolution of terrestrial environments of the Upper Silesian Keuper as biotopes of vertebrates” (N N307 11703) granted for Grzegorz Racki by the Ministry of Science and Higher Education. At least in the stratigraphic context, this ASGP thematic issue should be a conclusive heuristic stimulus for the more consistent studies, including vertebrate researches. Nevertheless, a prospect for understanding the mostly unresolved problems are outlined as well.
At least three widely separated bone-bearing intervals in the Upper Triassic succession of Upper Silesia, ranging in age from the Carnian to Rhaetian (i.e., in the interval of 25 Ma), are presented in papers of the Warsaw research group, led mostly by Jerzy Dzik and/or Grzegorz Niedźwiedzki. The stratigraphic arguments are reviewed for so far studied vertebrate localities, in particular for well-known middle Keuper sites at Krasiejów and Lisowice-Lipie Śląskie, to show that previously proposed age assignments are still inadequately documented and questionable. This unreliability is exemplified by evolving stratigraphic correlations of the fragmentary Silesian sections (10-18.5 m thick) with informal subsurface units from central-western Poland and with the German standard succession, ultimately not corroborated by comparison with the composite reference succession of the Upper Silesian Keuper, including new Woźniki K1 and Patoka 1 well profiles (ca. 260 m thick). Based on a multidisciplinary stratigraphic study covering consistent litho-, bio-, climato- and chemostratigraphic premises, focused on the relatively complete regional reference section, two bone bed levels only are recognized in the Patoka Marly Mudstone-Sandstone Member (= Steinmergelkeuper) of the Grabowa Formation, not very different in age (Classopollis meyeriana Palynozone; probably IVb Subzone): (1) localized Krasiejów bone breccia level (early Norian in age) in the Opole region, and (2) far more widely distributed Lisowice bone-bearing level (middle Norian) in a vast alluvial plain (braided to anastomosing river system) during the Eo-Cimmerian tectonic-pluvial episode. As a consequence of the principal uncertainties and controversies in Upper Triassic terrestrial stratigraphy, this is still a somewhat preliminary inference. Typical fluvial physical skeletal concentrations of combined hydraulic/sedimentologic type are common in the Upper Silesian Fossil-Lagerstätten, even if a conservation element is probably important as well.
Mass accumulations of vertebrate fossils in the tetrapod "graveyard" at Krasiejów near Opole, SW Poland, occur in a vast lacustrine marly claystone horizon and claystone lenses of various extent within fluviatile cross-laminated mudstone. These fossil assemblages do not differ from each other in taxonomic composition, but the proportions of aquatic to land animals are dramatically different. We attempted to separate these two components of the assemblages to restore the original composition of the biota. The lacustrine biocoenosis component of Krasiejów includes characean algae, various molluscs and arthropods, ganoid and dipnoan fishes, the phytosaur Paleorhinus, and the temnospondyl amphibian Metoposaurus as the most common vertebrate. The capitosaurid labyrinthodont Cyclotosaurus probably occupied the lake shore. The inland vertebrate community was dominated by the herbivorous aetosaur Stagonolepis and the small herbivorous dinosaur Silesaurus, which probably were the prey for the rauisuchian Teratosaurus. The geological age of the Krasiejów strata can be determined, although with a rather low resolution, based on position of various members of its fauna in their evolutionary lineages. Biostratigraphic and sequence stratigraphy evidence may improve the precision of this dating. The strata seem to correspond with the upper part of the Weser Formation in Germany, believed to be of Late Carnian age.
Charcoals from the Upper Triassic vertebrate-bearing clays of the Zawiercie area (Upper Silesia, S-Poland) were analyzed using petrographic methods, to reconstruct burning temperatures as well as taphonomic processes. SEM and reflected light microscopy show excellent preservation of charcoals most probable connected with early diagenetic permineralization by calcite. The charcoal was assigned to three morphotypes, probably corresponding to three different fossil taxa. Fusinite reflectance data suggest, that the highest temperature reached above 600 °C (fusinite reflectance of 3.59%), what counterparts to the lower limit crown fire temperature. The values for most of the samples are lower (ca. 1% to 2.5%) what is typical for surface fires. In many cases fusinite reflectance values depends on the measured zone within the sample. Such zonation formed due to charring temperature differences. In zones remote from the potential fire source, reflectance values gradually decreases. It implies that calculation of fire temperatures based on average fusinite reflectance values might be too far-reaching simplification. Occurrence of fungal hyphae within the charcoal supports the interpretation of a predomination of surface fire, consuming dead twigs and stems. The low content of micro-charcoals in charcoal-bearing rocks as well as roanded to sub-roanded shapes of large specimens indicates that they were transported after burning, deposited away from the burning area, and finally early diagenetic mineralization.
Age assignments of Triassic tetrapod fossils can be achieved by direct reference to a scheme of Triassic land-vertebrate faunachrons (LVFs) that correlates Triassic tetrapod fossil assemblages to each other based solely on the tetrapod fossils. Correlation of Triassic tetrapod assemblages to the standard global chronostratigraphic scale (SGCS, the “marine timescale”) is a separate cross correlation between the vertebrate biochronology and marine biochronology that usually relies on other data (e. g., palynostratigraphy, magnetostratigraphy, radioisoto-pic ages) to be completed. Late Triassic tetrapod fossils in southern Poland are found at two stratigraphic positions, the Krasiejów and Lisowice levels. The tetrapod assemblage of the Krasiejów level is assigned to the early Adamanian LVF based primarily on the stratigraphic overlap of the phytosaur Parasuchus with the Adamanian index aetosaur Stagonolepis. The amphibians Cyclotosaurus and Gerrothorax, a Proterochersis-like turtle and the aetosaur Paratypothorax from the Lisowice level indicate it is assignable to the Revueltian LVF. Cross correlations to the SGCS are less definitive, but suggest that the Krasiejów level is late Carnian and the Lisowice level is early/middle Norian. However, this correlation of the Krasiejów level is confounded by disagreements over correlation of the marine Carnian-Norian boundary to nonmarine strata. Indeed, the possibility that the Krasiejów tetrapods fill a gap in the early Norian record of tetrapods merits consideration. Such difficulties emphasize the value of correlattng tetrapod assemblages to each other using a land-vertebrate biostratigraphy/biochronology, instead of immediately attempting the more problematic correlation to the SGCS.
The results ofthe palynostratigraphical studies presented in this paper come from five boreholes Patoka 1, Czarny Las, Woźniki Śląskie K1, Kobylarz 1 and Poręba as well as from four outcrops at Lipie Śląskie, Patoka, Zawiercie and Poręba, in Upper Silesia (southern Poland). The palynostratigraphical zonation presented by Orłowska-Zwolińska (1983) for the epicontinental Upper Triassic of Poland was applied. The palynomorph spectra are marked by different preservation states, combined with the frequent occurrence of reworked specimens, probably even from Palaeozoic strata. The spore-pollen assemblage recognized in the “Chrzanów Formation” belongs to the early Carnian verrucata Subzone of the palynological longdonensis Zone. The spectrum from the Stuttgart Formation represents the Carnian astigmosus Zone. Spec tra in the Patoka Marly Mudstone-Sandstone Member (Grabowa Mudstone-Carbonate Formation), with the Lisowice bone-bearing horizon, represent the middle and late Norian meyeriana b Subzone. The Rhaetian age of the bone-bearing succession in the Lisowice-Lipie Śląskie clay-pit suggested in the literature was not confirmed. The age of assemblages from the “Polomia Formation”, which overlies the Patoka Member, was not determined, owing to the poor state of miospore preservation. Moreover, three types of palynofacies were recognized as being characteristic for a fluvial channel (1), a flood plain (2), and lacustrine and playa environments (3) as well as for an undetermined milieu. Type 1 was found in the deposits of the Stuttgart Formation, the Patoka Member and the “Polomia Formation”, type 2 in the Patoka Member and the “Polomia Formation”, type 3 in the “Chrzanow Formation”, the Stuttgart Formation and the Patoka Member.
Premise of research. Abundant fossil and molecular evidence suggests that all extant conifer families wereestablished by the Early Cretaceous. However, the recognition and understanding of the lineages that lead to theevolution of these extant families remain incomplete. Late Paleozoic to Early Mesozoic Voltziales conifers-also known as transitional conifers, usually with multilobed ovuliferous scales-have been hypothesized tobe among the stem lineages of modern crown conifers. This article describes an exquisitely preserved voltzianseed cone from the Aptian-Albian of Mongolia that introduces new taxonomic diversity and morphological datainto the complex pattern of conifer evolution.Methodology. Bulk lignite samples collected from the Tevshiin Govi locality were disaggregated in water,cleaned with hydrochloric and hydrofluoric acids, washed, and dried in air. Fossils were examined using LM,SEM, and X-ray microtomography.Pivotal results. Krassilovia mongolica gen. et sp. nov. has seed cones with helically arranged, imbricated,and tightly interlocked bract-scale complexes. Each mature bract-scale complex consists of an inconspicuousbract partially fused to the stalk of a five-lobed scale. Three of the lobes are distal (always pointing away fromthe cone base), while the other two are proximal (always pointing toward the cone base). Up to five invertedwinged seeds are present on the adaxial side of the ovuliferous scales. A systematic review of Late Paleozoic to Early Cretaceous multilobed ovuliferous scales-together with a morphological cladistic analysis-supportsthe placement of the Mongolian material in a new genus within the voltzian clade of the Voltziales.Conclusions. Krassilovia mongolica provides evidence of the additional diversity of extinct voltzian conifersand shows that some persisted to inhabit forest-moor swamp environments in eastern Asia during the Early Cretaceous. The new fossil taxon also shows novel morphological adaptations of the bract-scale complexesand the cone (i.e., imbrication and interlocking) for the protection of the ovules/seeds that are broadlyconcurrent with the appearance of new insect and other animal feeding strategies.
Seed cones (Compsostrobus neotericus gen. et sp. nov.), pollen cones, and vegetative remains of coniferophytes occur in Upper Triassic rocks of the Deep River Basin (Pekin Formation) of Central North Carolina. Seed cones have spatulate ovuliferous scales, each with two ovules and subtended by an elongated bract with an attenuate tip. Cuticle of seed cones resembles that of leaves on vegetative axes. Slender leaves are borne along two sides of the axis. Pollen cones have helically arranged microsporophylls, each with two abaxial sporangia bearing pollen grains of the Alisporites type. Seed cones, pollen cones, and vegetative remains suggest a coniferophyte very modern in aspect.
Describes the results of mineralogical, petrographical and paleobotanical investigations of petrified fragments of tree logs, found in Poreba near Zawiercie. The fossils in question represent fragments of the coniferous, Rhaetico-Liassic trees, silicified under the conditions of the tropical climate. The fragments have been determined as Araucarioxylon sp. The fossils of Poreba, despite difficulties in definition of their genus affinity, deserve a detailed description being rare findings in Mesozoic rocks of Poland. -from Authors
This book provides up-to-date coverage of fossil plants from Precambrian life to flowering plants, including fungi and algae. It begins with a discussion of geologic time, how organisms are preserved in the rock record, and how organisms are studied and interpreted and takes the student through all the relevant uses and interpretations of fossil plant. With new chapters on additional flowering plant families, paleoecology and the structure of ancient plant communities, fossil plants as proxy records for paleoclimate, new methodologies used in phylogenetic reconstruction and the addition of new fossil plant discoveries since 1993, this book provides the most comprehensive account of the geologic history and evolution of microbes, algae, fungi, and plants through time. * Major revision of a 1993 classic reference * Lavishly illustrated with 1800 images and user friendly for use by paleobotanists, biologists, geologists and other related scientists * Includes an expanded glossary with an extensive up-to-date bibliography and a comprehensive index * Provides extensive coverage of fungi and other microbes, and major groups of land plants both living and extinct.
Six male cones from the "Lower Keuper" of Bromsgrove (Worcestershire) referred by Wills (1910) to Voltzia heterophylla and assigned by Townrow (1962) to Masculostrobus willsi are reinvestigated. These ones resemble the male cones of the Voltzia sandstone (Buntsandstein) of the Vosges (France) which have been assigned to Masculostrobus rhomboidalis (Grauvogel-Stamm 1969). A comparison of their in situ pollen grains is the subject of this paper.