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Evidence of widespread fungal attack on Upper Triassic trees in the southwestern USA
Abstract
Certain Upper Triassic tree trunks in the southwestern U.S.A. show evidence of damage similar to that caused in tree trunks at the present day by pocket rot fungi such as Polyporus amarus Hedgc. and Heterobasidion annosum (Fries) Bref. The damaged trunks occur in a relatively thin stratum at the same horizon over a wide area in the Petrified Forest National Park, Arizona and elsewhere in the southwestern United States duting the Late Triassic. If the peculiar pattern of damage was caused, as it seems, by a pathogenic fungus then it can be compared in scale with the extensive destruction of Ulmus trees by Dutch Elm Disease [Ophiostoma (Ceratocystis) ulmi (Buisman)] in recent years in Europe and North America.
... No rigorous attempt was made to identify the trees fossilized here, as taxonomic classification may require specific description of the tracheid structure, which is difficult in partially degraded wood. Nonetheless, the wood most likely can be attributed to Araucarioxylon arizonicum Knowlton, the most common woody taxon in Chinle strata, as other genera are known primarily only from a single horizon in the Chinle, the Black Forest Bed in the Painted Desert Member (Creber and Ash, 1990), which occurs well above the Sonsela Member (Heckert and Lucas, 2002). ...
... Correlation of the beds within the Sonsela Member from PFNP to the study area, approximately 220 km to the northwest, is somewhat problematic. Nevertheless, we feel confident that the material we describe was collected within the uppermost strata of the Sonsela Member, and therefore occurs stratigraphically above the horizon containing the material described by Creber and Ash (1990). ...
... These are longitudinal channels that are similar in size and morphology to the axial channels we find. Creber and Ash (1990) attributed these features (of both scales) to the degrading actions of pathogenic fungi, a pocket rot fungus in particular, such as Polyporus amarus. They noted that fungus does not abrade the tracheids, as an invertebrate burrower might, and that it travels through the heartwood of the tree. ...
Permineralized wood from the middle of the Upper Triassic Petrified Forest Formation in Arizona, in strata correlative with the Sonsela Member, displays various forms of heartwood degradation. Pitting of the wood exhibits two primary morphologies. Elongated cavities are up to several mm long, parallel to the wood grain and may merge to form channels that extend longitudinally through the wood for lengths of several cm. Circular to elliptical cavities that cross-cut the wood grain are up to several mm wide. Both forms of degradation are inter-preted as the result of pre-burial biotic activity. The pitting appears to be caused by pathogenic fungi (white rot and white pocket rot in particular) that degraded and removed tracheids in the secondary xylem. The size and shape of some of the features, such as the longitudinal channels, may be consistent with some forms of arthropod burrowing, but the weight of the evidence, in particular the lack of frass, suggests that these also resulted from fungal decay.
... 7) agreed with Roadifer that the Rainbow Forest sandstone represents an upper tongue of the Sonsela Sandstone bed and was " part of the same fluvial system " . Ash (1987) and Creber and Ash (1990) recognized the Rainbow Forest sandstone as a separate unit below the Sonsela. Murry (1990) described an informal sandstone on Camp Butte northeast of Blue Mesa and interpreted it as a possible lateral equivalent to the Rainbow Forest sandstone. ...
... ttern on the surfaces similar to that seen in the siliceous horizon found in assemblage D. Other locations exhibit increasing size and abundance of siliceous rhizoliths culminating in a nearly continuous horizon, similar to a silcrete horizon in morphology (Klappa, 1980; Wright and Tucker, 1991), or simply a horizon with a silcrete-like morphology. Creber and Ash (1990) also described silicified whole logs, branches and roots with a " rope-like " texture within this horizon. Facies assemblage E is interpreted as floodplain, avulsion , and channel deposits. Large channels (>3m thick) commonly show evidence of high sinuosity by abundant and welldefined lateral accretion surfaces (Thomas et al., 1987; Bri ...
... A range in floodplain positions relative to the channel, from proximal to distal, is interpreted from lateral relationships and the high variability in pedogenic features, including slickenside abundance and size, matrix and mottling color and carbonate abundance, suggesting variation in both duration of pedogenesis and drainage conditions (e.g., Bown and Kraus, 1987; Kraus, 1987; Bown and Kraus, 1993; Aslan and Autin, 1998). The siliceous horizon was interpreted by Creber and Ash (1990) as an interval of increased silification due to increased pore space from a period of widespread fungal attack on plant material . These authors also commented on the potential regional stratigraphic utility this horizon. ...
INTRODUCTION PETRIFIED FOREST National Park (PEFO) is a focal point for studies of Upper Triassic terrestrial strata in the American Southwest. Despite numerous studies of plant and vertebrate remains, stratigraphic assessment of the Chinle Formation has lagged behind. Previous biostratigraphic study has indicated a change in the fauna and flora in the middle of the PEFO sec-tion, surrounding the level of the Sonsela Sandstone (Long and Padian, 1986; Litwin et al., 1991; Lucas and Hunt, 1993; Long and Murry, 1995; Murry and Kirby, 2002). However, detailed lithostratigraphic studies have not concentrated on this important interval, despite their importance for any biostrati-graphic framework. Lithostratigraphic study of the PEFO region, south of the Navajo and Hopi Reservations, began in earnest with Cooley (1957; 1958; 1959) and Akers et al. (1958). Stewart et al. (1972a) produced an overview of the regional lithostratigraphy, but did not focus on the PEFO section. Billingsley et al. (1985) produced the first full geologic map of PEFO at a 1:50,000 scale and Billingsley (1985) published a companion explanation of PEFO stratigraphy. Billingsley's (1985) and Billingsley et al.'s (1985) work largely followed the nomenclature of previous workers, and was the standard for the stratigraphy of PEFO that was followed by most workers for the next 15-20 years with only local additions and revision (Ash, 1987; Ash, 1992; Therrien and Fastovsky, 2000; Hasiotis et al., 2001). During the 1990s views regarding the stratigraphy of the Chinle Formation, and thus PEFO, split into two main phi-losophies. One philosophy continued along the lines of Stewart et al. (1972b) and Billingsley, Breed, and Ash (Billingsley, 1985; Billingsley et al., 1985) maintaining informal regional lithologic unit designations, and emphasizing regional correlations be-tween units (e.g Lucas et al., 1999) who incorporated all Upper Triassic terrestrial strata of western North America into the Chinle Group and either for-malized existing local nomenclature into broad members, aban-doned existing stratigraphic nomenclature or changed the strati-graphic rank of units. Recently, Heckert and Lucas (2002b) expanded the Sonsela Member (=Sonsela Sandstone bed) within PEFO by introducing a tripartite subdivision. The new revision is similar to what is observed at the type section of the Sonsela (Akers et al., 1958) and incorporates some observations seen by pre-vious workers that indicated that the Sonsela within PEFO consists of several sandstone beds (Cooley, 1957; Roadifer, 1966). However, the stratigraphy as proposed by Heckert and Lucas (2002b) includes correlations of subunits contradic-tory to mapping and does not provide a sound foundation for the recognition of units on a regional level. Regional recogni-tion of the Sonsela has been complicated in the past by subjec-tive interpretation of isolated sandstone bodies that may or may not be restricted to the Sonsela interval throughout the region. The present study was initiated independently of Heckert and Lucas (2002b) and aims to provide a consistent ABSTRACT— The Sonsela Sandstone bed in Petrified Forest National Park is here revised with specific lithologic criteria. It is raised in rank to Member status because of distinct lithologies that differ from other members of the Chinle Formation and regional distribution. Informal stratigraphic nomenclature of a tripartite subdivision of the Sonsela Member is proposed that closely mimics past accepted nomenclature to preserve utility of terms and usage. The lowermost subdivision is the Rainbow Forest beds, which comprise a sequence of closely spaced, laterally continuous, multistoried sandstone and conglomerate lenses and minor mudstone lenses. The Jim Camp Wash beds overlie the Rainbow Forest beds with a gradational and locally intertonguing contact. The Jim Camp Wash beds are recognized by their roughly equal percentages of sandstone and mudstone, variable mudstone features, and abundance of small (<3 m) and large (>3 m) ribbon and thin (<2 m) sheet sandstone bodies. The Jim Camp Wash beds grade into the overlying Flattops One bed composed of multistoried sandstone and conglomerate lenses forming a broad, sheet-like body with prevalent internal scours. The term "lower" Petrified Forest Member is abandoned in the vicinity of PEFO in favor of the term Blue Mesa Member to reflect the distinct lithologies found below the Sonsela Member in a north-south outcrop belt from westernmost New Mexico and northeastern Arizona to just north of the Arizona-Utah border. The Petrified Forest Member is here restricted in the vicinity of PEFO to the red mudstone-dominated sequence found between the Sonsela Member and the Owl Rock Member. 17 18 MUSEUM OF NORTHERN ARIZONA BULLETIN NO. 62 framework for the Sonsela interval on which to base other studies. Lithologic criteria are given to recognize the Sonsela and its three subunits, and justify the status of the Sonsela as a member of the Chinle Formation. The lithostratigraphic frame-work presented here provides a background for recognition of the Sonsela on a regional level, as well as locally within the PEFO area.
... As a result, there are some well-documented examples of fungal associations with land plants (Taylor et al. 2015). However, although showing some damaged wood, more or less well-preserved, the fossils do not always reveal hyphae or reproductive structures of the putative fungal pathogens ä Fig. 15 (continued) specimen, section K5459b1 showing one mature root with stele (S), the cavity of the former cortex now filled with sand (CC), and exodermis layer (arrow), between exodermis layer, and the periphery of the boring tiny lower-order roots are visible, scale bar ¼ 5 mm; (c) detail of Fig. 15b showing the triarch stele with protoxylem (arrows), exarch metaxylem (MX), secondary xylem (SX), and extraxylary tissue (ET) of the root, scale bar ¼ 500 μm; (d) detail of a juvenile root with stele (S), cortex cavity (CC) filled by sand, and exodermis layer (arrow) (note an additional side root (R) with stele and cortex cavity), section K5459b1, scale bar ¼ 500 μm; (e) juvenile root with stele (S), parenchymatous cortex (C) with lacunae, and exoderm layer (arrow), section K5459c, scale bar ¼ 500 μm (Creber and Ash 1990). Already Daugherty (1941) had attributed pockets of necrotic tissue in conifers from the Petrified Forest National Park, Arizona, USA, to fungal activity and introduced the fossil fungus Polyporites wardii. ...
... The expanding replacement in our material could also point to early diagenetic fixation of the bodies, in contrast to shrinkage that may have affected the surrounding wood. Similar patterns have been reported by Creber and Ash (1990) for petrified conifer trees from the Upper Triassic of the Southwestern United States. In the end, we cannot provide strong evidence whether the decomposition pattern found in our wood was caused by microorganisms or by any inorganic influence during its taphonomic pathway. ...
... Decay patterns consistent with white rot and white pocket rot have been observed in Glossopteris root wood (Vertebraria) and stem wood (Australoxylon) from the Permian of Antarctica (Stubblefield and Taylor, 1986;Weaver et al., 1997;Harper et al. 2017), and in Araucarioxylon wood from Triassic strata of the same region (Stubblefield and Taylor, 1986). The symptoms of wood-rotting fungi have also been described from numerous trunks of Late Triassic trees from the Petrified Forest of Arizona (Chinle Formation; Creber and Ash, 1990). Erasmus (1976, figs 1, 7) also illustrated, but did not describe, white and soft rots in an araucarian wood from the Cretaceous of Natal. ...
... The limited fossil record of saproxyly persists through the Triassic. During that interval, terrestrial saproxylic communities probably remained similar to those of the Permian in being dominated by fungi and arthropods, including oribatid mites and insects (Walker, 1938;Linck, 1949;Depape and Doubinger, 1963;Zhou and Zhang, 1989;Creber and Ash 1990;Kellogg and Taylor, 2004;Gnaedinger et al., 2012;Tanner and Lucas, 2013;Strullu-Derrien et al, 2012;McLoughlin and Strullu-Derrien, 2015;Gnaedinger and Zavattieri, 2020). Surprizingly, these communities are generally less well known than their Permian equivalents, most having lower apparent diversity, although they seemingly diversified somewhat during the Late Triassic (Kelber, 2007). ...
Organisms that colonize wood are subject to a taphonomic tragedy-the richer and more diverse they become, the greater the deterioration of the host wood and the less likely such communities are to be fossilized. Moreover, palaeobotanical studies of fossil wood usually focus on the plant tissue, neglecting the evidence of parasitic, saproxylic, and other contained organisms. Such a case involved a relatively well-known fossil wood assemblage from the Santonian (Late Cretaceous, ca 84 Ma) of southeast Africa. In a set of 150 thin sections of silicified wood stored in the Senckenburg Museum for more than half a century, we discovered evidence of a diverse biotic community comprising bacteria, fungi, nematodes , several types of arthropods, and marine bivalves. These body fossils and traces, together with growth-ring features, fossil log size and shape, and the distribution of glauconite, facilitated interpretation of the multi-stage evolution of a wood-hosted biocoenosis of unprecedented diversity. This record is unique for the Mesozoic and is of importance for understanding the taphonomic pathways to preservation and the evolution and diversification of saproxylic and other wood-hosted communities in terrestrial and marine settings. Ó
... As a result, there are some well-documented examples of fungal associations with land plants (Taylor et al. 2015). However, although showing some damaged wood, more or less well-preserved, the fossils do not always reveal hyphae or reproductive structures of the putative fungal pathogens (Creber and Ash 1990). Already Daugherty (1941) had attributed pockets of necrotic tissue in conifers from the Petrified Forest National Park, Arizona, USA, to fungal activity and introduced the fossil fungus Polyporites wardii. ...
... The expanding replacement in our material could also point to early diagenetic fixation of the bodies, in contrast to shrinkage that may have affected the surrounding wood. Similar patterns have been reported by Creber and Ash (1990) for petrified conifer trees from the Upper Triassic of the Southwestern United States. In the end, we cannot provide strong evidence whether the decomposition pattern found in our wood was caused by microorganisms or by any inorganic influence during its taphonomic pathway. ...
While knowledge about the paleobiology of plants has increased considerably, knowledge of the various interactions between trees, climbers, epiphytes, and other coexisting organisms has unfortunately remained limited. Besides morphology, anatomy, reproduction of plants, or even composition and distribution of their communities in space and time, only a thorough study of ecological aspects, the interrelatedness of all organisms, will lead to an all-encompassing
understanding of ancient forests. In this contribution, we examine diverse interactions between plants, animals, and putative microorganisms from Permian rocks of northern Tocantins state, central-north Brazil. The plant fossils derive from sedimentary strata of the Parnaíba Basin and are anatomically preserved as siliceous petrifactions. As major representatives of tropical seasonally influenced environments, these plants grew on mineral substrates. Their remnants comprise stems, branches, and roots of tree ferns, sphenophytes, and gymnosperms. Close-fitting or densely interwoven among each other, the plant organs and their various connections to other organisms contribute to the knowledge of multifaceted synecological relationships within the Northern Tocantins Fossil Forest. In some cases, however, the patchy fossil record must be treated with caution. Considering the identification of decay patterns in fossil wood, we are reliant on comparisons with modern plants because organic remains mostly lack in the petrifactions. Such comparisons are centered by the compartmentalization concept
focusing on patterns of wood reaction or partial decomposition in living trees, which mainly result from injuries or defense against the spread of pathogens. Though still consisting of a few glimpses into a usually hidden microworld, the unexpected diversity of organism interactions provides new insight into structure, function, and variety of riparian environments in low-latitude Southern Hemisphere regions.
... As a result, there are some well-documented examples of fungal associations with land plants (Taylor et al. 2015). However, although showing some damaged wood, more or less well-preserved, the fossils do not always reveal hyphae or reproductive structures of the putative fungal pathogens (Creber and Ash 1990). Already Daugherty (1941) had attributed pockets of necrotic tissue in conifers from the Petrified Forest National Park, Arizona, USA, to fungal activity and introduced the fossil fungus Polyporites wardii. ...
... The expanding replacement in our material could also point to early diagenetic fixation of the bodies, in contrast to shrinkage that may have affected the surrounding wood. Similar patterns have been reported by Creber and Ash (1990) for petrified conifer trees from the Upper Triassic of the Southwestern United States. In the end, we cannot provide strong evidence whether the decomposition pattern found in our wood was caused by microorganisms or by any inorganic influence during its taphonomic pathway. ...
This book will cover the entire evolutionary history that the terrestrial plants have recorded in Brazilian sedimentary rocks, ranging from the first vestiges of terrestrial environments colonization about 400 million years ago, until reaching the eve of the present time, when the current vegetation formations were organizing to reach their current distribution in modern biomes. At present Brazil is home to the world’s greatest plant biodiversity and we aim to offer here an opportunity to appreciate how this floral biodiversity originated and developed in these lowlands of South America, through chapters elaborated by the best Brazilian and foreign experts who dedicate to elucidate the evolution of the ancient flora in this part of the planet.
... Various decay structures and basidiomycotous fungal hyphae were recognized in the wood, including erosion troughs, cavities and varyingly decayed tracheids that are consistent with those caused by extant white-rot fungi (Stubblefield et al., 1985;Taylor, 1986, 1988). Similar wood decay structures have been documented from the subsequent Permian-Carboniferous (Diéguez and López-Gómez, 2005;Wan et al., 2016Wan et al., , 2017b, Mesozoic (Creber and Ash, 1990;Feng et al., 2015b;Gnaedinger et al., 2015;Tian et al., 2018;Zhu et al., 2018) and Cenozoic (Pujana et al., 2009). In the wood herein described, pocket-like cavities occur irregularly in the secondary xylem, which are comparable to modern white-rot characters produced by basidiomycetous fungi (Schwarze, 2007). ...
... The tracheids that surround these pockets show various cell-wall alterations, including removal of the middle lamella, separation of S1 and S3 walls, and the thickened tracheid corners. Although no fungal hyphae or spores were recognized, the recognition of similar pocket structures have also been reported in the Late Triassic gymnosperms from North America (Creber and Ash, 1990), Jurassic conifers from Argentina (García Massini et al., 2012;Sagasti et al., 2019), Cretaceous conifers from Antarctica (Falcon-Lang et al., 2001) and Eocene podocarpaceous wood from Argentina (Pujana et al., 2009) that contributed to fungal white-rot. ...
An exceptionally well-preserved wood, Ningxiaites shitanjingensis sp. nov., is described from the uppermost Permian Sunjiagou Formation of Northwest China providing new evidence for plant diversity and palaeoecological features of the Cathaysian Flora. The new plant is characterised by a eustelic vascular system with thick pycnoxylic woody cylinder. The cylindrical pith is solid and parenchymatous, surrounded by numerous, discrete endarch primary xylem strands and monarch leaf traces. The wood is especially distinctive by the expanded rays at the pith periphery and the pitted tangential walls of ray cells. The tracheids in the primary xylem have annular, helical and scalariform thickenings from protoxylem to metaxylem. The secondary xylem comprises tracheids, rays and axial parenchyma cells. Tracheids in the secondary xylem possess both alternate and opposite bordered pits on the radial walls. Each cross-field displays 1–2 cupressoid bordered pits. Rays are mostly uniseriate, 1 to 14 cells high. Leaf traces originate individually, bifurcate or not during their horizontal extension through wood, form helical clusters. Two types of arthropod coprolites were recognised in a boring in the pith. The relatively smaller coprolites containing unidentifiable plant tissues are identical with those of oribatid mites, whereas the larger coprolites exhibiting undigested plant cells probably produced by ancient millipedes. In addition to the occurrences of evident fungal hyphae, white-rot pattern and tyloses in the delignificated wood. The current study demonstrates complex ecological relationships and trophic networks occurred during the Lopingian (late Permian).
... Fungi play a fundamental role as decomposers and recyclers of organic matter in terrestrial ecosystems, thereby maintaining the carbon cycle, essential to the continuation of life on Earth (Carlile et al. 2001, Dighton 2016. Lignicolous fungi are the main causes of wood decay within modern ecosystems because of their enzymatic ability to degrade lignin, hemicellulose and cellulose; the fossil record shows that this interaction is well represented deep into the geological past (Cridland 1962, Stubblefield et al. 1985, Stubblefield & Taylor 1986, Creber & Ash 1990, Taylor & Osborn 1992, Weaver et al. 1997, Cantrill & Poole 2005, Diéguez & López-Gómez 2005, Pujana et al., 2009, García Massini et al. 2012b, Tanner & Lucas 2013, Gnaedinger et al. 2015, McLoughlin & Bomfleur 2016, Harper et al. 2017; however, the scarcity of direct fossil evidence of this kind of plant-fungi interactions, and the low representation of the current spectrum of known patterns of wood rot in the geological record, make the fossil woods from the Río Leona Formation especially important for better understanding the interaction between these organisms and the palaeoecological significance of the various strategies used by fungi as decomposers of wood in ancient ecosystems. ...
... A remarkable example from the Jurassic of Antarctica demonstrates white-rot decay resulting from invasion by a pathogenic fungus that triggered the development of tyloses in its plant host as a barrier against decay (Harper et al. 2012). Another interesting example is the massive fungal decay described in various tree trunks from the Late Triassic of North America, which was compared with similar widespread fungal attacks in modern forests (Creber & Ash 1990). The current example from Patagonia supports the widespread presence in the geological record of white-rot fungi as carriers of wood decay either as saprotrophs or necrotrophs, which were critical for nutrient recycling through time. ...
Carlos Daniel Greppi, Juan L. García Massini, Roberto R. Pujana and Sergio A. Marenssi, May 2018. Fungal wood-decay strategies in Nothofagaceae woods from Miocene deposits in southern Patagonia, Argentina. Alcheringa xxx, xxx -xxx.
Decayed woods from the Miocene, Rio Leona Formation, Santa Cruz, Argentina having simultaneous decay patterns consistent with soft- and white rot characteristics are described. Samples were previously identified as Nothofagoxylon scalariforme. At low magnification, the permineralized woods appear mottled, with discoloured, degraded areas, scattered in apparently robust tissue, consistent with white-rot decay. At greater magnification, the woods reveal several micromorphological features, including differential decay of cellulose-rich cellular components that match soft-rot decay by extant ascomycetes and some basidiomycetes. In addition, decayed woods either appear differentially delignified or show simultaneous decay of all cellular components (lignin- and cellulose-rich), which are by-products of white-rot fungal decay. Additional anatomical characteristics of the decayed woods are consistent with a host response to the fungal attack. Co-occurrence of these two decay patterns suggests soft-rot decay and white-rot fungal decay. In addition, co-occurrence of all the decay features observed also suggests facultative soft rot by white-rot fungi, such as in some extant species that switch between these two types of decay strategies as a means to circumvent plant defences. These data indicate that fungi with soft-rot capacity for wood decay can be traced back to the early Miocene (ca 19 Ma). In addition, this report adds to the distribution and diversity of fungi in the geological record and underscores the ecological importance of wood as a preferred substrate for the association and interactions between fungi with different saprotrophic abilities, which have been fundamental for nutrient recycling in terrestrial ecosystems during the Cenozoic.
Carlos Daniel Greppi [greppi.carlos.d@gmail.com] Museo Argentino de Ciencias Naturales-CONICET, Av. Ángel Gallardo 470, (1405) Ciudad Autónoma de Buenos Aires, Argentina; Juan L. García Massini [massini112@yahoo.com.ar] Centro de Investigaciones Científicas y Transferencia Tecnológica de La Rioja (CRILAR), Provincia de La Rioja, UNLaR, SEGEMAR, UNCa, CONICET, Entre Ríos y Mendoza s/n (5301) Anillaco, La Rioja, Argentina; Roberto R. Pujana [rpujana@gmail.com] Museo Argentino de Ciencias Naturales-CONICET, Av. Ángel Gallardo 470, (1405) Ciudad Autónoma de Buenos Aires, Argentina; Sergio A. Marenssi [smarenssi@hotmail.com] IGEBA-CONICET, Departamento de Geología, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Intendente Guiraldes 2620, (1428) Buenos Aires, Argentina.
... Fungal activity and other kinds of microbial decay produce structures in wood and leaves that can be considered traces, though little attention has been devoted to them. Exceptions include Stubblefield and Taylor (1986), Creber and Ash (1990), Kelber (2007) (2013), and Taylor et al. (2015). As discussed below, much more attention has been paid to the study of microbial decay in bones. ...
... Some laterally continuous records of these ichnofacies also are well documented. For example, the Paleoscolytus ichnofacies has extensive records, such as the fungal traces in numerous Late Triassic fossil logs from Arizona (Creber and Ash, 1990) and the many prolific assemblages that show leaf damage amounting to hundreds/ thousands of fossil leaves in the Upper Cretaceous of the Western Interior (e. g., Labandeira et al., 2002). Bonebeds, particularly of Late Jurassic dinosaurs, provide aerially extensive assemblages assignable to the Cubiculum ichnofacies (e.g., Bader et al., 2009), and laterally continuous (formation-wide) assemblages of fossil bone from the Upper Cretaceous of Montana preserve diverse MFD (Lam et al., 2009). ...
Substantial ichnocoenoses of ichnofossil assemblages in nonmarine environments can be grouped into two, substrate-controlled ichnofacies. These are traces in plant tissues (mostly wood and leaves) and in bone. Plant tissues and bones provide distinctive, “hardground” substrates that are colonized and/or utilized in the nonmarine realm by a substantial diversity of organisms, mostly arthropods and microorganisms. The trace fossil record in plant substrates is diverse, extending back to the Devonian, and is dominated by borings in wood created for feeding, reproduction, and shelter and by feeding traces on leaves, including mines and galls. The Paleoscolytus ichnofacies named here includes nonmarine trace fossil ichnocoenoses dominated by feeding traces and borings of low to moderate ichnodiversity. Woody and foliar substrates typify the Paleoscolytus ichnofacies, and it is to some extent the subaerial counterpart of the previously named Teredolites ichnofacies. Bone provides a source for feeding, reproduction, and shelter for various arthropods; it is also a source of food for vertebrates, and it is modified in some settings by trampling and by human activities. Bone also undergoes bioerosion by microbial agents—bacteria, fungi and protozoans. The result is a diversity of traces preserved in fossil bone that encompass distinctive ichnofossil ichnocoenoses. The Cubiculum ichnofacies named here includes nonmarine and subaerial trace fossil ichnocoenoses dominated by macroscopic and microscopic borings produced by mobile feeders and the subordinate occurrence of damage by other bone utilizers, of generally low to moderate ichnodiversity and of high local abundance.
... As with the Martha's Butte beds, interbedded mudstones are rich in pedogenic carbonate nodules (Herrick, 1999;Woody, 2006;Martz and Parker, 2010). A distinctive reddish or orange horizon composed of silicified plant material occurs low in the unit (Creber and Ash, 1990;Martz and Parker, 2010;Driese et al., 2010), just below the level where pedogenic carbonate and unionids become abundant, and is associated with important faunal and floral overturns (Parker and Martz, 2011;Reichgelt et al., in press). Vertebrate fossils are extremely abundant. ...
... Sandstone, siltstone and mudstone; mudstones are purple and less commonly bluish or gray ( Figure 7B-C). In southern PEFO, exposures north of the Flattops mesas can locally be roughly divided into upper beds dominated by gray and reddish sandstone and conglomerate with minor purple and bluish mudstone, and lower beds dominated by purple mudstone interbedded with discontinuous gray and dull pinkish ribbon and sheet sandstones which are not usually very conglomeratic ( Figure 7B; Martz and Parker, 2010 (Creber and Ash, 1990;Woody, 2006;Martz and Parker, 2010). ...
The Upper Triassic Chinle Formation in Petrified Forest National Park represents some of the most intensively studied Upper Triassic strata in western North America. Five stratigraphic members are exposed within the park, from oldest to youngest: the Mesa Redondo, Blue Mesa, Sonsela, Petrified Forest, and Owl Rock Members. Despite numerous stratigraphic studies of the Chinle Formation and two attempts at mapping the park over the past sixty years, sandstone marker beds in the Sonsela Member at the north and south ends of the park were still poorly mapped and correlated. Studies in the years 2002 and 2006 claimed that two sandstones which previous workers had considered to lie at different stratigraphic levels (the Jasper Forest Bed and the Flattops One sandstones in the Martha's Butte beds) were actually correlative. This correlation resulted in a three-part division of the Sonsela Member and had a major impact on vertebrate biostratigraphy. In a recent attempt to resolve confusions regarding Chinle Formation lithostratigraphy and biostratigraphy, we have completely walked out lithologic contacts through most of the park. The resulting new geologic map, revised lithostratigraphic model, and associated data resolves the 2002 and 2006 miscorrelations by demonstrating that the Jasper Forest Bed capping Blue Mesa and Agate Mesa and Flattops One sandstones (Martha's Butte beds) are stratigraphically distinct, resulting in a thicker and more complex five-part model for the Sonsela Member, and considerably modifying the vertebrate biostratigraphy. New geologic mapping also resulted in a detailed lithostratigraphic framework for the northern park which has previously been poorly understood, and several important new marker beds, including a purple-gray bed that represents the base of the Owl Rock Member. The revised geologic map is an ArcGIS product that includes an updated lithostratigraphic model for the Chinle Formation, fossil localities, and hyperlinks to labeled photographs of measured sections. A pre-existing ArcGIS product created by workers at Northern Arizona University was used for the creation of this map and thus the final product includes some mapping, mostly Quaternary alluvium, dunes, and sandsheets, from that study. The cumulative effect of these revisions is to emphasize the importance of thoroughly exploring stratigraphic contacts, extensively documenting lithostratigraphic models, making georeferenced GIS maps, and accurately locating critical fossil localities. These methods and the new map make lithostratigraphic, biostratigraphic, and paleoecologic and paleoclimate models scientifically testable to future researchers at this classic Chinle Formation location.
... Later, Pujana et al. (2009) showed similar traces in a piece of Eocene wood from Argentina effectively associated to cellular decay produced by fungi. Previously, Creber and Ash (1990) have shown similar fungal attacks on Upper Triassic trees from the Chinle Formation (USA), but represented by elongated rods instead of pits. ...
... It may be an indicator of humid or subhumid terrestrial palaeoenvironments according to its fungal origin. Particularly when the fungal attack is widespread in entire forests as those of the Chinle Formation (Creber and Ash, 1990) or the Allen Formation presented herein. A. lignorum, being fungus traces produced in continental settings, has a very different paleoenvironmental meaning that A. xylobiontum. ...
... Auch werden bei der natürlichen Verkieselung die Einflüsse veränderter Durchlässigkeit für siliziumhaltige Fluida und veränderte pH-Werte diskutiert, wenn nach Ursachen für unterschiedliche Farben und Erhaltungszustände petrifizierter und permineralisierter Hölzer gesucht wird (z. B.Mustoe & Acosta 2016, Creber & Ash 1990, Rößler et al. 2021. Ein Beleg, dass durch den pilzlichen Stoffwechsel Lignin im Clover-Creek-Fossil bevorzugt abgebaut wurde, könnte beim Verständnis des noch immer nicht gänzlich geklärten Prozesses der Holzversteinerung helfen. ...
Zeigt die fossile Eiche Quercinium pliocaenicum makroskopische Anzeichen von Kernfäule durch Weißfäulepilze? Ausgangspunkt ist ein kurzer Überblick über den Metabolismus der Fäulnispilze unter Bevorzugung von Lignin oder Zellulose sowie die Fähigkeit der Bäume, sich in bedingtem Maße dagegen zu schützen (CODIT). Davon ausgehend werden Stücke einer 1895 in Clover Creek, Idaho, USA, gefundenen fossilen Eiche, Quercinium plio-caenicum, untersucht. Teile davon zeigen Verfärbungen, die in Vergleich zu rezenten Eichen zu der Annahme führen, dass zu Lebzeiten des Baumes Kernfäule durch Weißfäulepilze aufgetreten ist. Dadurch hätte sich das Verhältnis der Holzbestandteile Zellulose und Lignin verändert und den Prozess der Verkieselung möglicherweise beeinflusst. Die in den dunkel verfärbten Abschnitten wesentlich schlechtere Erhaltung der organischen Gewebe suggeriert die Holzzerstörung vor der Einkieselung, lässt jedoch keine Reste pilzlicher Mikroorganismen mehr erkennen. Die Untersuchung des Übergangs von unverfärbtem zu verfärbtem Kieselholz mittels mikro-Röntgen-fluoreszenz ergab Anreicherungen von Verbindungen des Aluminiums, Eisens, Kaliums, Kalziums, Mangans und Zinks. Letztere kommen in der Nähe von Rissen sowie in unregelmäßig umgrenzten Teilen des dunkel verfärbten Kieselholzes vor, sind jedoch nicht an die SiO 2-Matrix gebunden. Daher führen wir die Anreicherungen von Me-tallionen auf Vorgänge während oder nach der Fossilisation zurück. Der mutmaßliche pilzliche Holzabbau vor der Fossilisation lässt sich nur durch den stärkeren Zersetzungsgrad des Holzes in den braun gefärbten Arealen belegen. Abstract Does the fossil oak Quercinium pliocaenicum show macroscopic signs of heart rot caused by white-rot fungi? We begin with a brief overview of two groups of wood-decaying fungi and their ability to derive nutrition from degrading cell-wall components as well as the tree's limited ability to protect itself according to the CODIT model. Patterns exhibited by specimens of the fossil oak Quercinium pliocaenicum found along Clover Creek, Idaho, USA, in 1895 are assessed in light of damage caused by wood-decaying fungi and the CODIT model. We propose a hypothesis that differential patterns of color and damage preserved in the opalized wood can be attributed to the effect of white-rot fungi when the tree was still growing. Some white-rot fungi preferentially degrade lignin, which could have changed the ratio of cellulose to lignin in the secondary xylem thus probably affecting the silicification process. The much poorer preservation of organic tissues in the brown mineralized areas suggests wood destruction prior to silicification, but does not reveal remnants of fungal microorganisms. Examination of the transition from yellow to dark brown silicified wood by micro-X-ray fluorescence revealed elemental enrich-ments of aluminum, iron, potassium, calcium, manganese, and zinc. The latter accumulations occur in cracks and irregularly bounded portions of the brown mineralized areas, but are not linked to the SiO 2 matrix. Therefore, we attribute the accumulations of metal ions to processes during or after fossilization. The presumed fungal degradation of wood prior to fossilization can be evidenced only by the greater degree of wood decomposition in the brown-colored areas.
... At the late stage of decay, preferential lignin degradation leads to the decomposition of the middle lamella, and individual secondary tracheid walls being separated from each other (Schwarze, 2000). Similar wood decay structures have been widely documented from the Paleozoic (Diéguez and López-Gómez, 2005;Wan et al., 2017a), Mesozoic (Creber and Ash, 1990;Gnaedinger et al., 2015;Zhu et al., 2018;Xia et al., 2020), and Cenozoic (Pujana et al., 2009;Ruiz et al., 2020b). In the current fossil stem of Protophyllocladoxylon yiwuense, pocket-like structures irregularly occur in the secondary xylem (Fig. 6E), which are comparable to the decay structures caused by modern white-rot fungi. ...
A new permineralized stem, Protophyllocladoxylon yiwuense Gou et Feng sp. nov., is described from the Middle Jurassic Xishanyao Formation in Naomaohu Town of Yiwu County, Hami City, Xinjiang Uygur Autonomous Region, Northwest China. The stem is only preserved with secondary xylem, which is pycnoxylic and exclusively composed of tracheids and parenchymatous rays. The tracheids have uniseriate or biseriate, alternately arranged bordered pits on their radial walls, and uniseriate bordered pits on the tangential walls. The rays are uniseriate or partially biseriate, up to 34 cells high. There are one or two large, simple pits in each cross field. A branch trace horizontally passes through the secondary xylem showing anatomical structures identical to the secondary xylem. Analyses of the growth-ring width and mean sensitivity values indicate that P. yiwuense had long growing seasons with readily available water supplies and favorable paleoclimatic conditions. White-rot decay features are commonly recognized in the stem, including pocket-like cavities, complete removal of the middle lamellae, thickened corners and separation of the secondary walls of tracheids, as well as septa-like structures in the tracheid lumens due to the cell's reactions to fungal activity. Our study sheds new light into the plant diversity and paleoenvironmental and paleoecological conditions of the Middle Jurassic.
... Silicified woody trunks are playing an important role in the palaeobotanic (Falcon-Lang & Bashforth, 2004;Falcon-Lang & Scott, 2000;, palaeoecological (Creber & Ash, 1990;Feng, Schneider, Labandeira, Kretzschmar, & Röβler, 2015;Feng, Wang, & Liu, 2010;Feng, Wang, Rößler, Ślipi nski, & Labandeira, 2017;García Massini, Falaschi, & Zamuner, 2012;Wan, Yang, Liu, & Wang, 2016;Wan, Yang, & Wang, 2014;Wei, Gou, Yang, & Feng, 2019) and palaeoclimatic (Creber & Chaloner, 1984, 1985Creber & Francis, 1999;Falcon-Lang, 1999;Taylor & Taylor, 1993;Wan, Yang, He, Zhou, et al., 2017;Wan, Zhou, Yang, & Wang, 2016;Zhang, Wang, Liu, & Li, 2010) research. Their anatomical characteristics provide unique information for the systematics and physiology of gymnosperms and other plant groups (Falcon-Lang, Kurzawe, & Lucas, 2014;Wan, Yang, Tang, Liu & Wang, 2017), which are beneficial for realizing plant diversity in the geological periods. ...
A silicified trunk, Zhuotingoxylon liaoi Wan, Yang, Wang, Liu et Wang gen. et sp. nov., is described from the uppermost part of Guodikeng Formation in South Taodonggou section, Turpan–Hami Basin, Xinjiang Uygur Autonomous Region, northwestern China. It is characterized by a solid pith, endarch primary xylem and pycnoxylic wood. The pith is composed of parenchyma and sclereids. Radial walls of primary xylem tracheids have spiral and scalariform thickenings. Secondary xylem consists of thick‐walled tracheids and parenchymatous rays. Uniseriate rounded pits with oval apertures are distributed on radial tracheidal walls separately. Cell walls of rays are homogeneous and smooth. Rays are 1–10 cells high in tangential section. Cross‐field pits are cupressoid. There are 1–4 bordered pits with slit‐like to oval apertures in each cross‐field. Based on the anatomical features of the pith and xylems, it is proposed that the new stem has a coniferous affinity. The new fossil stem adds to the knowledge of vascular plant diversity close to the Permian–Triassic boundary.
... Despite the dense occurrence of cell wall fragments within these bands, there is no development of damaged pockets that are devoid of cells inside the bands. However, the damage pattern points to parasitic rather than saprophytic decay, which would show a more diffuse pattern of attack (Creber and Ash, 1990). ...
For the first time, this study describes the dynamics of white rot fungal decay in a petrified conifer branch with clear araucarian affinity from the late Aptian Crato Lagerstätte (Santana Formation, Araripe Basin, northeastern Brazil). High resolution optical microscopy was used to identify tridimensional chemical and anatomical evidence in different regions of the bark and xylem tissues of permineralized shoots, and results support the hypothesis that the host responded to disease that may have started when it was still alive. The wood decay pattern was strongly indicative of the selective decay by white rot. The general pattern of interaction is consistent with pathogenic rather than saprophytic fungal activity. Analysis of fungus–plant interactions associated with growth ring patterns imply intermittent periods of favorable temperature-moisture inputs that were crucial for fungal activity during the deposition of the Crato fossil Lagerstätte included in the Tropical Equatorial Hot arid belt.
... Thin-sections or polished sections through one of the hollows would help to assess their cause, but because this was the only tree found possessing hollows we did not section it for curational reasons. A thin section would have helped because bacterial-fungal activity leaves a typical trace in the wood (Creber & Ash 1990). ...
Fossil tree hollows are seldom described in the literature and can often be elusive to the field paleobotanist. However, these structures
may provide unique paleoecological, environmental and tree life history information that are essential for a more complete
understanding of ancient forests. A stump from the ‘late Permian’ (Wuchiapingian–Changhsingian) of the Mágoè Fossil Forest in
Mozambique (Tete Province) provides a rare example of fossilized tree hollows. These hollows were found near the base of the tree and
are subcircular in shape, ranging between ~1.3 and 3.5 cm in diameter. Although thirty-one trees were densely sampled (i.e. no fossil
trees were excluded from a given area, in our case ~2650m2) and inspected at the Mágoè Fossil Forest, only one (PPM2017-31) exhibited
tree hollows, highlighting the scarcity of these structures in this fossil forest. In modern forests tree hollows are more likely to be found
in old trees, likewise PPM2017-31 was among the largest trees found in the sample, suggesting this was an old tree. The subcircular
morphology of the tree hollows indicates they resulted from fungal/bacterial activity rather than from a fire.
... The geological record preserves a variety of examples of multitrophic interactions, which have provided insight into timing of appearance and type of associations between different groups of organisms in ancient ecosystems (Taylor et al., 2009(Taylor et al., , 2015Krings et al., 2016). In particular, fossil plants preserving evidence of interactions with fungi and xylophagous arthropods have been previously described from multiple localities (e.g., Müller-Stoll, 1936;Creber and Ash, 1990;Ibáñez and Zamuner, 1996;Harper et al., 2012;García Massini et al., 2012b;Klymiuk et al., 2015;Falcon-Lang et al., 2015). Peronosporomycetes have been found acting as pathogens in the periderm of Lepidodendron (Carboniferous of France; Dotzler et al., 2008) and in conifer embryos (Triassic of Antarctica), in which they occur associated with putative zygomycetes (Schwendemann et al., 2010). ...
Article available at: https://authors.elsevier.com/a/1Xwbc73NzuYCI
A permineralized araucarioid conifer stem, preserved in state of partial decay, is described from the Middle to Late Jurassic Chon-Aike Formation (Bahía Laura Group) at Laguna Flecha Negra, in the Deseado Massif, Santa Cruz Province, Argentina. The tree-stem, which comprises periderm to the secondary xylem, shows evidence of
interactions with xylophagous borers, fungi, actinomycetes and other fungal-like microorganisms. In the secondary phloem, it displays rows of traumatic resin ducts, which host abundant mycelium comprising very thin hyphae differentiated into spore-producing structures comparable to actinomycete bacteria and spherical fungallike resting structures. In the cortical and phloem regions, there are cell groups with structural changes, which appear variably degraded by fungi. The secondary xylem displays decay of cellulose and lignified components, at various spatial scales, comparable with fungal decay patterns in modern wood. Further septate fungal filaments, sometimes bearing clamp connections are associated with decayed regions. The degraded secondary xylem shows circular to oval borings filled with frass containing degraded tracheids, comparable to those bored by different wood boring beetles. Development of traumatic resin ducts in the phloem could have been triggered by penetration of the living stem by these xylophagous arthropods. The association of the different components present suggests pathogenic and saprotrophic interactions promoting decomposition of the stem, and represents a rarely-documented example of multitrophic biological interactions involving several different biological groups in Jurassic terrestrial paleoecosystems.
... Thin-sections or polished sections through one of the hollows would help to assess their cause, but because this was the only tree found possessing hollows we did not section it for curational reasons. A thin section would have helped because bacterial-fungal activity leaves a typical trace in the wood (Creber & Ash 1990). ...
Fossil tree hollows are seldom described in the literature and can often be elusive to the field paleobotanist. However, these structures may provide unique paleoecological, environmental, and tree life history information that are essential for a more complete understanding of ancient forests. A stump from the Wuchiapingian-Changhsingian of the Mágoè Fossil Forest in Mozambique (Tete Province) provides a rare example of fossilized tree hollows. These hollows were found near the base of the tree and are subcircular in shape, ranging between ~1.3 to 3.5 cm in diameter. Although thirty-one trees were densely sampled and inspected at the Mágoè Fossil Forest, only one (PPM2017-31) exhibited tree hollows, highlighting the scarcity of these structures in this fossil forest. In modern forests tree hollows are more likely to be found in old trees, likewise PPM2017-31 was among the largest trees found in the sample, suggesting this was an old tree. The subcircular morphology of the tree hollows indicates they resulted from fungal/bacterial activity rather than from a fire.
... By the Late Triassic (ca. 237-201 Ma), cambium engravings are associated with the grazing of the secondary xylem (wood) outer surface and periderm (bark) inner surface [11][12][13] . These borings of wider diameter give rise to irregular, smaller, side-branch tunnels 12,13 . ...
Beetles are the most diverse group of macroscopic organisms since the mid-Mesozoic. Much
of beetle speciosity is attributable to myriad life habits, particularly diverse-feeding strategies
involving interactions with plant substrates, such as wood. However, the life habits and early
evolution of wood-boring beetles remain shrouded in mystery from a limited fossil record.
Here we report new material from the upper Permian (Changhsingian Stage, ca. 254–252
million-years ago) of China documenting a microcosm of ecological associations involving a
polyphagan wood-borer consuming cambial and wood tissues of the conifer Ningxiaites
specialis. This earliest evidence for a component community of several trophically interacting
taxa is frozen in time by exceptional preservation. The combination of an entry tunnel
through bark, a cambium mother gallery, and up to 11 eggs placed in lateral niches—from
which emerge multi-instar larval tunnels that consume cambium, wood and bark—is ecologically
convergent with Early Cretaceous bark-beetle borings 120 million-years later.
... Most wood-borers fall into this category: woodpeckers tap trees in spring to obtain sap (e.g., Glutz von Blotzheim, 1980;Schwenke, 1986), shipworms do not only protect themselves in the substrate but also digest it (e.g., Turner, 1966), moss mites bore in various woods leaving characteristic microcoprolites (e.g., Brongniart, 1877;Labandeira et al., 1997), termites and gribbles digest cellulose by gnawing corridors (e.g., Becker et al., 1957;Hartnack, 1943;Krishna et al., 2013;Menzies, 1957;Tappen, 1994), various Coleoptera bore into wood as larvae (e.g., Bostrychidae, Brenthidae, Buprestidae, Cerambycidae, Cupedidae, Curculionidae, Dermestidae, Lucanidae, Lyctidae, Lymexylonidae, Oedemeridae, Psoidae, Ptinidae, Scolytidae, Serropalpidae) or adults (Anobiidae, Anthribidae, Bostrychidae, Ostomidae, Platypodidae, Scolytidae), mostly leaving traces that are characteristic of a species or at least a genus (e.g., Ratzeburg, 1837;Escherich, 1923;Vit e, 1952;Schwenke, 1974;Hickin and Edwards, 1975;Brauns, 1991). Although ecosystemarily important, boring bioerosion of fungi in wood is rather rarely preserved, but it results in typical structures (e.g., Creber and Ash, 1990;Genise et al., 2012). In the case of bone, these may be tineid moths (e.g., Davis and Robinson, 1999;Huchet, 2014), darkling beetles (Holden et al., 2013), the siboglinid annelid Osedax (e.g., Goffredi et al., 2005), fungi (e.g., M€ agdefrau, 1937;Jans, 2008), and bacteria (e.g., Deming et al., 1997;Amano and Little, 2005;Kaim et al., 2008;Jans, 2008). ...
Bone is a substrate for bioerosion at equal rank with xylic and lithic substrates. Accordingly, borings in bone have to be identified in an analogous way to other ichnogenera coined for one type of substrate. In due course, the new ichnogenera Osteichnus n. igen. and Clavichnus n. igen. are established within the new ichnofamily Osteichnidae. Gastrochaenolites and Trypanites are here restricted to lithic substrates, and Asthenopodichnium only occurs in xylic substrates. Only with this approach, ichnotaxobases of trace fossils in bone are identical to those in other hard substrates. Cuniculichnus variabilis n. igen. n. isp. is introduced for variably shaped pits to tunnels bored into bone by beetle (arguably dermestid) larvae; its ethological character is close to a pupichnion.
... Examples of fungal decay in Late Permian wood include Dadoxylon Endlicher specimens from the Iberian Ranges of Spain that exhibit selective delignification and cell wall separation (Diéguez & López-Gómez 2005). Symptoms of wood rot have also been described from the Upper Triassic Petrified Forest of Arizona, but it was not possible to determine whether the infections were parasitic or saprotrophic because no bona fide fungal remains were detected (Creber & Ash 1990). Reports of fungal remains associated with wood decay symptoms include Xenoxylon yunnanensis Feng, a gymnosperm wood from the Middle Jurassic of China that displays decay symptoms in the form of micro-pockets most similar to extant white pocket rot, with associated fungal hyphae that possess well-defined clamp connections (Feng et al. 2015). ...
Evidence of fungal decay is frequently encountered in silicified wood. However,
studies focusing on fossil fungal wood degradation remain rare. A characteristic
pattern of degradation and decay symptoms congruent with present-day white
pocket rot occur in Late Permian silicified glossopteridalean stem and root
wood (Australoxylon sp.) from Skaar Ridge, Antarctica. Co-occurring with the
decay symptoms are fungal hyphae with clamp connections. Hyphae usually
progress through the pit apertures, but some may also penetrate tracheid walls.
The individual wall layers in some of the infected tracheids are separated from
each other, apparently forming appositions. Small, opaque bodies (?arthropod
coprolites) occur in some of the decay pockets. The abundance of infected
specimens among the silicified woods from Skaar Ridge suggests that white
pocket rot fungi were important decomposers in late Paleozoic high-latitude
forest ecosystems.
... The present material shows diffusedly and evenly decayed areas all over the wood rather than the precisely defined pockets of rot. According to Creber and Ash (1990), the damaged wood attacked by saprophytic white-pocket rot fungi would present similar distribution and dimension of decayed areas. Therefore, although it is difficult to absolutely evaluate the interactions between fungi and Septomedullopitys szei, the fungi occurred in current fossil wood are more likely to be saprophytic. ...
Several lines of evidence of plant-arthropod and plant-fungus interactions are documented from the Wuchiapingian Wutonggou low-order cycle (approximate equivalent to the Wutonggou Formation) in Tarlong valley, southern Bogda Mountains, Xinjiang Uygur Autonomous Region, northwestern China. Fossil wood, Septomedullopitys szei Wan, Yang et Wang, contains differentially-damaged areas. Spindle-shaped pockets in the fossil wood occurring in the secondary xylem are commonly free of organic remains. They are comparable in appearance to modern white-pocket rot caused by fungi. The tracheid walls around the decomposed areas are degraded from the middle lamellae to outer layers. Abundant branching and septate fungal hyphae in the decayed areas, ray parenchyma and tracheid lumina indicate that fungi are responsible for the wood decay. These fungi are partially regarded as basidiomycetes because of the occurrence of clamp connections. According to the characteristic damages they caused to the host, ascomycetes are also viable candidates of the fungi because large parts of hyphae are without certain clamp connections. The other damaged excavations are branched and maze-like borings and galleries, which are filled with abundant fungal hyphae, cellular debris and spheroidal to ovoidal, dark-colored coprolites, ranging from 26 to 128 μm in diameter. The size, shape, and surface texture of these coprolites indicates that the coprolites are the feces of ancient oribatid mites. The fungal hyphae, coprolites, and degraded excavations in the pith of the late Permian wood suggest that wood-rotting and -boring were not limited to the xylem.
... Most of the wood likely can be attributed to the fossil species widely known as Araucarioxylon arizonicum Knowlton (Jones et al., 2002), the most common woody taxon in Chinle strata, although many authors now refer the genus Araucarioxylon to Agathoxylon (Philippe, 2011;Philippe, 2015;Byers et al., 2014;Degani-Schmidt and Guerra-Sommer, 2016). Other genera are known primarily only from a single horizon in the Chinle, the Black Forest Bed in the Painted Desert Member (Creber and Ash, 1990), which occurs above the Sonsela Member (Heckert and Lucas, 2002) and is not correlative with any of the fusain occurrences described here. ...
We document the occurrence of Upper Triassic fusain in northern Arizona, southern Utah and northern New Mexico in latest Carnian(?) to Norian-age alluvial strata of the Monitor Butte Formation and the Sonsela and Painted Desert members of the Petrified Forest Formation (Chinle Group). The fusain, identified by standard techniques of macroscopic and microscopic morphology, and resistance to chemical oxidation, is present at multiple stratigraphic horizons at several locations that are approximately correlative, although separated by several hundred km. The morphology of this fossil charcoal includes large, partially charred silicified to coalified logs, completely charred smaller limbs, charcoal fragments reworked in fluvial deposits, and particulate fusain disseminated in sediments. The apparent rarity of fusain in Triassic strata previously was cited as evidence for relatively low levels (compared to modern) of atmospheric oxygen and the consequent infrequent occurrence of wildfire during this time interval. However, our findings demonstrate a significantly greater abundance of fusain in Upper Triassic strata than previously acknowledged. Therefore, we suggest that wildfire was not an unusual occurrence during the Late Triassic. This conclusion supports recent experimental studies and geochemical models that indicate atmospheric oxygen levels at or above modern levels during the Late Triassic.
... Evidence of white rot pockets have also been reported in Late Triassic gymnosperms from North America (Creber and Ash, 1990), Jurassic conifers from Argentina (García Massini et al., 2012) and middle Cretaceous conifers from Antarctica . However, no fungal remains such as hyphae or spores have been recognised in these wood specimens. ...
... In their gross morphology, these structures are similar to some extant white-pocket rots caused by fungi (Blanchette, 1992). Similar damage also is present in Permian and Triassic conifer trunks from Antarctica (Stubblefield and Taylor, 1986) and North America (Creber and Ash, 1990), Jurassic conifers from Argentina (García Massini et al., 2012), and Cretaceous conifers from Antarctica (Falcon-Lang et al., 2001). ...
... In the fossil record, direct proof of fungal virulence can only be obtained from structurally preserved plant fossils showing traces of fungal attack. Permian and Triassic records are very rare, but silicifi ed wood has provided strong evidence that pathogenic wood-rotting fungi could have been well established at the time of the Late Permian biosphere crisis, while patterns of wood decay were similar to those found today (Stubblefi eld and Taylor, 1986;Creber and Ash, 1990). So far, infection by soilborne Rhizoctonia-like pathogens has only been recognized in silicifi ed rhizomes of Eocene plants (Lepage et al., 1994). ...
Throughout the world, latest Permian records of organic-walled
microfossils are characterized by the common presence of remains of filamentous organisms, usually referred to the palynomorph genus Reduviasporonites. Although generally regarded as indicators of global ecological crisis, fundamental controversy still exists over the biological and ecological identity of the remains. Both fungal and algal affinities have been proposed. We seek to resolve this enigma by demonstrating close morphological similarity of the microfossils to resting structures (monilioid hyphae, sclerotia) of Rhizoctonia, a mod- ern complex of soil-borne filamentous fungi that includes ubiquitous plant pathogens. By analogy with present-day forest decline, these findings suggest that fungal virulence may have been a significant contributing factor to widespread devastation of arboreal vegetation at the close of the Permian Period.
... Wood remains may also record the pattern of damage caused by bark beetles and fungal attack. For example, Creber and Ash (1990), on the basis of welldefined pockets of decay, were able to attribute damage on Upper Triassic trees in the south-western USA to a pathogenic pocket rot fungus rather than a saprophyte. ...
The temperate and boreal forests of Europe and North America have been subject to repeated pathogen (fungal disease and phytophagus insect) outbreaks over the last 100 years. Palaeoecology can, potentially, offer a long‐term perspective on such disturbance episodes, providing information on their triggers, frequency and impact. Mid‐Holocene declines in Tsuga and Ulmus pollen dominate the Quaternary literature on forest pathogens, yet the role of pathogens, and even the presence of pathogenic fungal diseases, during these events has yet to be established. Pathogen‐focused research strategies, informed by the sequence of events documented in modern outbreaks, and undertaken at high temporal resolution using a multi‐proxy approach, are required. It is argued that forest pathogens are likely to have been significant agents of past vegetation change, even in cases where climate change was the primary stress factor. Copyright © 2013 John Wiley & Sons, Ltd.
... Widespread fungal infections in fossil forest stands in environments subjected to recurrent ecological disturbance have been suggested to debilitate plants, possibly making them increasingly more susceptible to pathogenic fungi (opportunistic pathogens) (Creber and Ash 1990;Falcon-Lang and Cantrill 2001). Regarding this, it has been indicated that the amount of resources used by extant pathogenic fungi to disperse their propagules decreases with increasingly higher density and geographic proximity of their host(s) (Damgaard 1999). ...
Several stages of the life cycle of an endoparasitic fungus of the Chytridiomycota, here assigned to the extant genus Synchtrium, are described as the new species per- micus from silicified plant remains from the Late Permian (~250 Ma) of Antarctica. The thallus of Synchtrium permicus is holocarpic and monocentric and consists of thick- walled resting sporangia, thin-walled sporangia, and zoospores in different stages of development. A life cycle is hypothesized from the range of developmental stages. The life cycle begins when zoospores encyst on the host cell surface, subsequently giving rise to thin-walled sporangia with motile spores. Some zoospores (haploid) function as isogamous gametes that may fuse to produce resting sporangia (diploid). Roots, leaves, and stems of plants are among the tissues infected. Host response to infection includes hypertrophy. Morphological and developmental patterns suggest similarities with the Synchytriaceae (Chytridiales), particularly with Synchytrium. Previous records of chytridiomycetes are known from the Devonian Rhynie Chert and from the Carbonif- erous and the Eocene of the northern hemisphere; this report is the first on chytridio- mycetes from the Permian.
... Seedling Mesozoic. Interestingly, Creber and Ash (1990) report decay inoculation experiments (Worrall et al. 1983; Cobb et al. in fossilized wood from Upper Triassic trees in the south- 1989) and field sampling studies employing isozyme analyses western United States that resembles decay caused by Oligo- (Otrosina et al. 1992) provided evidence of host specificity in porus amarus (Hedgc.) Gilbn. ...
Isolates of Heterobasidion annosum (Fr.) Bref. representing North American S and P and European S, P, and F intersterility groups were subjected to isozyme analysis. European S, P, and F groups had more variability than the North American S and P groups in expected hterozygosity, number of alleles per locus, and percent polymorphic loci. In contrast with the North American S and P groups, the European intersterility groups could not be distinguished from each other on the basis of individual isozyme loci, although significant differences in allele frequencies exist between European S and P groups. This suggests that evolution proceeded at different rates in the intersterility groups, or intersterility barriers appeared later in the European populations relative to the North American populations of H. annosum. Changes in climate and host species associations during the Tertiary may have been a major factor in evolution of H. annosum intersterility groups. Key words: allozymes, forest tree hosts, playnological events, evolutionary relationships, Hymenomycetes, root disease.
... Considering the widespread occurrence of fungal remains within fossil woods from other periods (e.g. Creber and Ash, 1990;Scott, 2000), such an interpretation of fungal remains in Late Permian wood does not appear to be convincing. ...
... Examples in recent times are the chestnut (Castanea) decline in North America (Anderson, 1974;Hepting, 1974;Anagnostakis, 1995), the decline in elm (Ulmus) populations in Europe and North America due to outbreaks of Dutch elm disease (Karnosky, 1979;Stipes and Campana, 1981;Brasier, 1991) and the current reduction of butternut/walnut (Juglans) (Nicholls, 1979;Prey and Kuntz, 1982) and dogwood (Cornus) (Daughtrey et al., 1988; populations in eastern North America. There is evidence that tree populations have been subject to similar infections during prehistoric times, including the distant past (Creber and Ash, 1990). During the mid-across the Ulmus decline (Graumlich, 1993) have increased the understanding of ecological changes taking place at particular sites (Garbett, 1981;Peglar, 1993). ...
The decline in Ulmus pollen frequencies that occurred ca. 5000 14C years ago before present (BP) is a key biostratigraphic marker horizon in northwest European pollen diagrams, although its causes are still a subject of debate. To investigate this event further, fungal spore analyses were carried out across the Ulmus decline at Moel y Gerddi, north Wales, United Kingdom. The Ulmus decline was in three phases, with a primary decline with low cereal and Rumex pollen records as the only agricultural indicators. This was followed by a more significant decline, with general forest opening and the grassland/pasture indicator Plantago lanceolata. A third, less significant, decline was again accompanied by cereal-type pollen. Tree pollen frequencies subsequently recovered, with cereal-type pollen remaining well represented. The fungal data recorded woodland taxa and a background level of the obligate dung fungus Sporormiella, a proxy for local herbivore abundance. Sporormiella frequencies increased greatly after the main Ulmus decline, around which there were high percentages of the ascospores of the wood rot fungus Kretzschmaria deusta. The neoecology of Kretzschmaria deusta, and the behavior of its spore curve suggests the colonisation of local populations of already severely wounded trees. At this site Ulmus, and perhaps Tilia were infected at a time of markedly increased inferred herbivore concentrations. Neolithic farming techniques could have provided mortally wounded trees while enhancing livestock grazing, although the role of disease must also be considered. Elevated Kretzschmaria deusta values may be of ecological significance in the interpretation of the causes and nature of the Ulmus decline and similar forest disturbance events, and the indicator role of this fungus in forest paleoecology requires further study.
Several samples of fossilized wood (charcoal) were collected in the Papayita archaeological site, in coastal Ecuador. This carbonized material was encountered inside a layer of volcanic ash that sealed the site. The ash-sized tephra was produced by a sub-Plinian eruption from the Guagua Pichincha volcano contempora-neous with the late Valdivia phases during the Formative Period. Each of the samples was sectioned into 10 to 15 subsamples and examined under a Scanning Electron Microscope (SEM), producing high-resolution images with a large depth of field where the anatomical structures and their geochemical composition were vividly discernible. Each sample corresponds to organic matter of vegetable origin, that is, carbonized wood in the form of small rocks, whose appearance is that of carbonized woody tree trunks and or branches. We were able to observe vascular structures, specifically bundles of xylem. It was possible to conclude that these tracheids underwent a physicochemical transformation typical of petrification processes, leaving the molds intact. This allowed us to determine structural elements that support the identification of the group of plants to which these samples belong, through the methodology of comparison of the anatomical components of current species. The fossilized wood structures are three-dimensional and present characteristics that correspond to the group of higher plants, Gymnosperms, of the Podocarpaceae group. Among them, quadrangular tracheids, circular hole-shaped pits in the vascular system, and absent resin canals stand out. Central to the analysis is the presence of transverse parenchyma, which can be ascertained to correspond to vegetation from climates that are temperate or cold.
Uranium–lead (U–Pb) geochronology was conducted by laser ablation – inductively coupled plasma mass spectrometry (LA-ICPMS) on 7175 detrital zircon grains from 29 samples from the Coconino Sandstone, Moenkopi Formation, and Chinle Formation. These samples were recovered from ∼ 520 m of drill core that was acquired during the Colorado Plateau Coring Project (CPCP), located in Petrified Forest National Park (Arizona).
A sample from the lower Permian Coconino Sandstone yields a broad distribution of Proterozoic and Paleozoic ages that are consistent with derivation from the Appalachian and Ouachita orogens, with little input from local basement or Ancestral Rocky Mountain sources. Four samples from the Holbrook Member of the Moenkopi Formation yield a different set of Precambrian and Paleozoic age groups, indicating derivation from the Ouachita orogen, the East Mexico arc, and the Permo-Triassic arc built along the Cordilleran margin.
A total of 23 samples from the Chinle Formation contain variable proportions of Proterozoic and Paleozoic zircon grains but are dominated by Late Triassic grains. LA-ICPMS ages of these grains belong to five main groups that correspond to the Mesa Redondo Member, Blue Mesa Member and lower part of the Sonsela Member, upper part of the Sonsela Member, middle part of the Petrified Forest Member, and upper part of the Petrified Forest Member. The ages of pre-Triassic grains also correspond to these chronostratigraphic units and are interpreted to reflect varying contributions from the Appalachian orogen to the east, Ouachita orogen to the southeast, Precambrian basement exposed in the ancestral Mogollon Highlands to the south, East Mexico arc, and Permian–Triassic arc built along the southern Cordilleran margin. Triassic grains in each chronostratigraphic unit also have distinct U and thorium (Th) concentrations, which are interpreted to reflect temporal changes in the chemistry of arc magmatism.
Comparison of our LA-ICPMS ages with available chemical abrasion thermal ionization mass spectrometry (CA-TIMS) ages and new magnetostratigraphic data provides new insights into the depositional history of the Chinle Formation, as well as methods utilized to determine depositional ages of fluvial strata. For parts of the Chinle Formation that are dominated by fine-grained clastic strata (e.g., mudstone and siltstone), such as the Blue Mesa Member and Petrified Forest Member, all three chronometers agree (to within ∼ 1 Myr), and robust depositional chronologies have been determined. In contrast, for stratigraphic intervals dominated by coarse-grained clastic strata (e.g., sandstone), such as most of the Sonsela Member, the three chronologic records disagree due to recycling of older zircon grains and variable dilution of syn-depositional-age grains. This results in LA-ICPMS ages that significantly predate deposition and CA-TIMS ages that range between the other two chronometers. These complications challenge attempts to establish a well-defined chronostratigraphic age model for the Chinle Formation.
Palynoflorules containing sparse but regularly occurring chitinous-walled fungal, probably ascomycete, spores have been obtained from silty limestone nodules in the Jurassic Coon Hollow Formation in the Wallowa Terrane in Hells Canyon, Idaho. The fungal spores are associated with moderately abundant embryophytic spores and pollen that suggest late Early Jurassic to early Middle Jurassic age, which agrees with dating provided by marine animals statigraphically just above. Two new species of the fossil fungal spore form-genus Diporicellaesporites, D. idahoensis and D. serratulus, and one new species of form-genus Fractisporonites, F. pittsburgensis, are described. -Authors
Silicified fossil woods were collected from an alluvial deposit in the Allier valley, Massif Central, France. These woods, dated as Oligocene, are identified as Quercoxylon lecointrei Gazeau & Koeniguer, a fossil species close to the extant Mediterranean holm oak (Quercus ilex L.).Thespecimens studied show numerous individual cavities, each 0.5-1.3 × 0.2-0.5 cm in size, but these generally do not affect multiseriate rays. Despite the lack of fungal remains, the decay pattern is comparable in appearance with that of presentday pocket-rot caused by Basidiomycetes, and particularly by Xylobolus frustulatus (Pers.: Fr.) Boidin. Indications of possible host response to fungal attack include abundant tyloses in vessels and dark substances filling the cells. ©2012 E. Schweizerbartsche Verlagsbuchhandlung, Stuttgart, Germany.
This paper contains a description of the only known non-banded bark in the geologic record. The bark is attached to a branch of the Araucarioxylon arizonicum tree that was found recently in the Late Triassic Chinle Formation (∼220 Mya) in Arizona. In the fossil the vascular cylinder is almost totally enclosed in rough bark 2-11 mm in thickness. The inner bark is less than 1 mm thick and comprises a very narrow zone of presumed phloem containing infrequent fibers. The outer bark varies in thickness up to 10 mm, and consists of curving lenses of sequent periderm free of both dilation tissue and resin canals but is quite unlike the banded bark of Araucaria heterophylla. Wide zones of very thin-walled rectangular parenchyma cells arranged in radial files and groups of thin-walled spongy parenchyma, and small clusters as well as solitary sclerenchyma occur in the rhytidome. The vascular cambium contains 1-4 fusiform cells per radial file. Uniseriate rays are common in the secondary xylem and extend to the cambial zone. Small (1 mm in diameter), simple tunnels in the bark and wood indicate that the branch was attacked by phytophagous insects. A possible petrified beetle larva similar in form and size to Anobium is associated with these tunnels.
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.
Mammalian assemblages in the Bembridge Limestone Formation of late Eocene age, Headon Hill, Isle of Wight, England, indicate habitats ranging from open woodland to closed forest. Distinctive 'lower' and 'upper' mammalian faunas reflect different faunal provinces, probably in response to climatic fluctuations that foreshadowed the terminal Eocene event. In order to improve our understanding of these patterns, we have examined a Variety of other palaeoenvironmental indicators from this section. These include paylnological organic matter (POM), plant macrofossils, non-mammalian faunas, organic geochemistry and stable isotopes. The evidence shows that the depositional setting was a tranquil, shallow, freshwater lake, with a brief lagoonal interval However, evidence for habitats surrounding the lake is contradictory, emphasizing the necessity for multidisciplinary approaches to palaeoenvironmental reconstruction. The mammal faunas give unequivocal evidence for woodland or forest, yet, apart from some of the land snails, there is no other indication of the presence of trees. Furthermore, according to the mammalian evidence the lake was bordered by distinctive vegetation at different times, with closed forest/woodland during marl deposition and open woodland during black mud deposition, but there are no parallel fluctuations in other biotic elements.
High-precision geochronology provides unprecedented insights into the depositional history of the Upper Triassic Chinle Formation of the Colorado Plateau, as well as its paleoenvironmental and paleobiological records. The Chinle succession exposed in the Petrified Forest National Park (PEFO) and vicinity, Arizona, includes two large-scale alluvial composite sequences. Although each composite sequence fines upward, the upper composite sequence is more dominated by coarser-grained deposits. Petrographic analysis of sandstone lithic content indicates an upward decrease in the proportion of volcanic rock fragments in each composite sequence. Paleocurrent indicators in the lower composite sequence suggest a variable paleoflow direction, whereas northward paleoflow dominated the upper composite sequence. The change in paleoflow appears to coincide with a reorganization of alluvial depositional processes and associated source terranes, and precedes a rapid acceleration in basin subsidence. Climate proxy records from paleosol geochemistry indicate a gradual shift from humid to dry conditions across the transition between the lower and upper composite sequences and the Adamanian-Revueltian biotic turnover. Composite-sequence depositional reorganization, climatic shift and biologic turnover, in turn, appear to coincide with episodes of magmatism recorded in Triassic granitoid plutons presently exposed in southern California. Taken collectively, these observations suggest that the Late Triassic depositional, climatic, and ecologic history at PEFO may be related to emergence of the incipient Cordilleran magmatic arc along the convergent western margin of Pangea. A new U-Pb date for the lower part of the Chinle Formation suggests that most or all of the formation was deposited in the Norian Stage.
Over the past 50 years the most enigmatic feature of pollen diagrams from northwest Europe has been the mid-Holocene 'elm decline', and there has been much speculation as to the origin(s) and cause(s) of this event. A total of 150 radiocarbon dates from 139 sites spanning the elm decline in Britain and Ireland have been collated and scrutinized. Statistical analyses on 138 dates show that the event has a mean date of 5036 14C yr BP with a standard deviation of ± 247. Calibration of the dates and combining the sum probabilities yielded a range spanning 6347-5281 cal yr BP (1σ), covering 1066 years. The start of the elm decline event lies between 6343 and 6307 cal yr BP (1σ), a period of 36 years, indicating that the onset was rapid. The end of the event lies between 5290 and 5420 cal yr BP (1σ), a period of 130 years. The probability distribution indicates that the elm decline was a uniform phased event across the British Isles. It appears that the elm decline can be explained to a large extent by the outbreak of disease. However, recent research on palaeoclimatic change and the nature of the transition from the Mesolithic to Neolithic in the British Isles suggests that both climatic change and human activities were implicated. It was probably the interplay between these factors, rather than any in isolation, that catalyzed the widespread, catastrophic decline of elm populations during the mid-Holocene.
The biodiversity and terrestrial ecology of the Late Albian Triton Point Formation (Fossil Bluff Group), Alexander Island, Antarctica is analysed to improve our understanding of polar biomes during the mid-Cretaceous thermal optimum. This formation was deposited on a high-latitude (75°S) floodplain and consists of two facies associations, a lower braided alluvial plain unit and an upper coastal meander-belt unit. Analysis of fossil plants in well exposed palaeosols reveals the existence of spatially complex plant communities. Braidplains supported patchy, low-density (91 trees/ha) stands of podocarp and taxodioid conifers on floodbasin substrates, and conifer-cycadophyte-fern-angiosperm thickets in riparian settings. Coastal meander-belts supported medium density (568 trees/ha) podocarp-araucarian conifer forests on mature floodbasin soils, and fern-angiosperm-ginkgo thickets in riparian settings. Growth-ring analysis indicates plants experienced stressful growing conditions on the braidplain characterized by high-frequency flood events, but more favourable growing conditions on the coastal plain. Additional vegetation disturbances were caused by arthropod-fungal attack, frost and wildfire. In terms of structure, composition, ecology and productivity these predominantly evergreen, broad-leafed conifer forests bear similarities to the extant temperate rainforests of New Zealand.
Four new ichnogenera and six new ichnospecies are described from permineralized plant remains from an unnamed Upper Cretaceous formation, of northern Patagonia in Argentina. This association of traces is the second ichnocenosis described for permineralized wood. Traces are interpreted as insect borings in wood and fruits. Cycalichnus garciorum n. ichnosp. is the first trace fossil recorded in a cycad stem and the oldest known evidence of a termite nest for the southern hemisphere. Stipitichnus koppae n. ichnosp. is the first trace fossil recorded in a palm stem. Xylonichnus trypetus n. ichnosp. is interpreted as borings of beetles in conifer wood. Carporichnus maximus n. ichnosp., C. bertheorum n. ichnosp., and C. minimus n. ichnosp. represent the first ichnocenosis related with a single species of fruit, and the only known Mesozoic borings in fruits. An historical overview of insect traces in permineralized wood shows that most were described ignoring the ichnological aspects, and this resulted in the absence of a taxonomic classification. In this paper, the main ichnotaxobases to describe insect traces in petrified wood are proposed for the first time.
Recent stratigraphic revisions of the Upper Triassic Chinle Formation of Petrified Forest National Park, in conjunction with precise and accurate documentation of fossil tetrapod occurrences, clarified the local biostratigraphy, with regional and global implications. A significant overlap between Adamanian and Revueltian faunas is rejected, as is the validity of the Lamyan sub-land vertebrate faunachron. The Adamanian–Revueltian boundary can be precisely placed within the lower Jim Camp Wash beds of the Sonsela Member and thus does not occur at the hypothesised Tr-4 unconformity. This mid-Norian faunal turnover, may coincide with a floral turnover, based on palynology studies and also on sedimentological evidence of increasing aridity. Available age constraints bracketing the turnover horizon are consistent with the age of the Manicouagan impact event. The rise of dinosaurs in western North America did not correspond to the Adamanian–Revueltian transition, and overall dinosauromorph diversity seems to have remained at a constant level across it. The paucity of detailed Late Triassic vertebrate biostratigraphic data and radioisotopic dates makes it currently impossible to either support or reject the existence of globally synchronous Late Triassic extinctions for tetrapods.
Although fungi have a long geologic history, many aspects regarding their origins and subsequent evolution remain impossible to document from the fossil record. As heterotrophs, fungi must interact with other organisms, and it is here that the fossil record can provide an important source of biological and paleoecological information about fungal interactions. Saprophytic, parasitic and biotrophic interactions among fungi and other organisms are ancient; examples of these interrelationships are discussed as they relate to the establishment and evolution of the biological and physical paleoecosystem.
Fossil evidence of terrestrial vascular plant life and terrestrial arthropods exists from the Silurian. Fossil evidence suggests progressive interaction between the two groups through the later Palaeozoic and Mesozoic. In this paper we present data, particularly from plant fossils, concerning several interactions: feeding, shelter, transport and reproduction. Evidence of arthropod feeding includes eaten leaves, borings in plant tissues, wound reaction and leaf mining as well as gut contents and coprolites from the arthropods themselves. We trace the changes in leaf eating behaviour from continuous marginal feeding, common in the Palaeozoic and early Mesozoic to the more abundant interrupted-marginal and non-marginal feeding behaviour on Cretaceous angiosperm leaves. This change may reflect the evolution of chemical defence strategies by the plants but also may reflect the evolution of different mouthpart design in new insect groups. Leaf mines and leaf galls, although known from the Upper Carboniferous, only become common in the Cretaceous, coinciding with the evolution of several new insect groups and plants. Wood boring is recorded, for the first time, from the Lower Carboniferous and becomes common from the Upper Carboniferous. Data from coprolites suggest that spore feeding preceded leaf feeding. Experiments using pteridophytes and living arthropods indicate that some spores remain viable after passing through the gut and hence this feeding habit may have also been advantageous to some early plants for propagule transport. We conclude that there is much evidence in the fossil record suggesting plant-arthropod interaction, but many more observations are required before detailed interpretations concerning coevolution can be made.
Recent revisions to the Sonsela Member of the Chinle Formation in Petrified Forest National Park have presented a three-part lithostratigraphic model based on unconventional correlations of sandstone beds. As a vertebrate faunal transition is recorded within this stratigraphic interval, these correlations, and the purported existence of a depositional hiatus (the Tr-4 unconformity) at about the same level, must be carefully re-examined.
Our investigations demonstrate the neglected necessity of walking out contacts and mapping when constructing lithostratigraphic models, and providing UTM coordinates and labeled photographs for all measured sections. We correct correlation errors within the Sonsela Member, demonstrate that there are multiple Flattops One sandstones, all of which are higher than the traditional Sonsela sandstone bed, that the Sonsela sandstone bed and Rainbow Forest Bed are equivalent, that the Rainbow Forest Bed is higher than the sandstones at the base of Blue Mesa and Agate Mesa, that strata formerly assigned to the Jim Camp Wash beds occur at two stratigraphic levels, and that there are multiple persistent silcrete horizons within the Sonsela Member.
We present a revised five-part model for the Sonsela Member. The units from lowest to highest are: the Camp Butte beds, Lot's Wife beds, Jasper Forest bed (the Sonsela sandstone)/Rainbow Forest Bed, Jim Camp Wash beds, and Martha's Butte beds (including the Flattops One sandstones). Although there are numerous degradational/aggradational cycles within the Chinle Formation, a single unconformable horizon within or at the base of the Sonsela Member that can be traced across the entire western United States (the "Tr-4 unconformity") probably does not exist. The shift from relatively humid and poorly-drained to arid and well-drained climatic conditions began during deposition of the Sonsela Member (low in the Jim Camp Wash beds), well after the Carnian-Norian transition.
Plant pathogens cause mortality and reduce fecundity of individual plants, drive host population dynamics, and affect the structure and composition of natural plant communities. Pathogens are responsible for both numerical changes in host populations and evolutionary changes through selection for resistant genotypes. Linking such ecological and evolutionary dynamics has been the focus of a growing body of literature on the effects of plant diseases in natural ecosystems. A guiding principle is the importance of understanding the spatial and temporal scales at which plants and pathogens interact. This review summarizes the effects of diseases on populations of wild plants, focusing in particular on the mediation of plant competition and succession, the maintenance of plant species diversity, as well as the process of rapid evolutionary changes in host-pathogen symbioses.
Gymnospermous secondary xylem degraded by wood-decaying fungi was examined in silicified fossils from Antarctica. Araucarioxylon-type wood from the Triassic and specimens of Vertebraria from the Permian demonstrate similar patterns of decay. Specimens are characterized by irregularly shaped areas lacking cells and are infected with branched, septate hyphae with clamp connections. The decay in these fossils is comparable in appearance to present-day rots caused by basidiomycetes. Two patterns of decay are evident: (1) A wall component, presumably lignin, is removed from the wall and middle lamella of infected tracheids, leaving a considerably thinner cellulose framework; and (2) the primary and secondary walls typically separate, and all wall layers are progressively reduced in thickness and eventually removed. The middle lamella, particularly where it is thickest in the corners between cells, persists longer than other layers. Indications of host response to fungal attack include the production of possible wall appositions. Evidence of wood decay occurs as early as the Upper Devonian, and all reported cases from the fossil record are similar to those found today.
summaryThe fossil record is a long-neglected source of valuable information concerning the diversity, structure, evolution and activities of ancient fungi. However, interest in these organisms has increased in recent years as their importance in a wide variety of interactions with plants, animals and the geological environment has been recognized. Evidence from the fossil record now implicates fungi in such processes as the establishment of terrestrial plants, degradation of lignin in Devonian forests, parasitic relationships with plants and animals, and the development of ancient soils. Although fossil fungi remain difficult to study, their geological record is rich and rewarding. Recent advances in the study of fungi from the Palaeozoic and early Mesozoic (exclusive of palynological research) are presented in this review.
The sequence of changes observed microscopically that occurred in wood throughout successive stages of decay was studied in the sapwood of a hardwood, sweetgum (Liquidambar styraciflua L.), and of a softwood, southern pine (Pinus sp.). The fungi used were Polyporus versicolor L. ex Fr., a white-rot fungus, and Poria monticola Murr., a brown-rot fungus. Light microscopy, plus the techniques of polarization and ultraviolet-absorption microscopy, was used to make the observations on sections 4 microns thick cut from celloidin-embedded specimens. (Author)
Disorders caused by non-living agents diseases caused by living agents on a wide range of hosts diseases of spruce diseases of pine diseases of larch diseases of Douglas Fir diseases of minor forest confiers - abiss, cupressus and chamaecyparis, thuja and tsuga diseases of other confiers diseases of oak diseases of sweet chestnut diseases of beech diseases of ash, birch and alder diseases of sycamore and maple diseases of elm diseases of poplar diseases of willow diseases of wild and ornamental rosaceus trees diseases of other broadleaved trees decay fungi of broadleaved and coniferous trees.
The Upper Triassic Flora of Arizona The Oxford Encyclopedia of Trees of the World Pathology of Trees and Shrubs
- L H Daugherty
Daugherty, L.H., 1941. The Upper Triassic Flora of Arizona. Carnegie Inst. Washington PUN., 526. Hora, B. (Editor), 1981. The Oxford Encyclopedia of Trees of the World. Oxford Univ. Press, Oxford, 288 pp. Peace, T.R., 1962. Pathology of Trees and Shrubs. Oxford Univ. Press, Oxford, 753 pp.
Tansley Review No. 12: Recent advances in palaeomycology
- Stubblefield