No support for the emergence of lichens prior to the evolution of vascular plants

The early‐successional status of lichens in modern terrestrial ecosystems, together with the role lichen‐mediated weathering plays in the carbon cycle, have contributed to the long and widely held assumption that lichens occupied early terrestrial ecosystems prior to the evolution of vascular plants and drove global change during this time. Their poor preservation potential and the classification of ambiguous fossils as lichens or other fungal–algal associations have further reinforced this view. As unambiguous fossil data are lacking to demonstrate the presence of lichens prior to vascular plants, we utilize an alternate approach to assess their historic presence in early terrestrial ecosystems. Here, we analyze new time‐calibrated phylogenies of ascomycete fungi and chlorophytan algae, that intensively sample lineages with lichen symbionts. Age estimates for several interacting clades show broad congruence and demonstrate that fungal origins of lichenization postdate the earliest tracheophytes. Coupled with the absence of unambiguous fossil data, our work finds no support for lichens having mediated global change during the Neoproterozoic‐early Paleozoic prior to vascular plants. We conclude by discussing our findings in the context of Neoproterozoic‐Paleozoic terrestrial ecosystem evolution and the paleoecological context in which vascular plants evolved.     

The Oldest Caseid Synapsid from the Late Pennsylvanian of Kansas,and the Evolution of Herbivory in Terrestrial Vertebrates

The origin and early evolution of amniotes (fully terrestrial vertebrates) led to major changes in the structure and hierarchy of terrestrial ecosystems. The first appearance of herbivores played a pivotal role in this transformation. After an early bifurcation into Reptilia and Synapsida (including mammals) 315 Ma, synapsids dominated Paleozoic terrestrial vertebrate communities, with the herbivorous caseids representing the largest vertebrates on land. Eocasea martini gen. et sp. nov., a small carnivorous caseid from the Late Carboniferous, extends significantly the fossil record of Caseidae, and permits the first clade-based study of the origin and initial evolution of herbivory in terrestrial tetrapods. Our results demonstrate for the first time that large caseid herbivores evolved from small, non-herbivorous caseids. This pattern is mirrored by three other clades, documenting multiple, independent, but temporally staggered origins of herbivory and increase in body size among early terrestrial tetrapods, leading to patterns consistent with modern terrestrial ecosystem.     

Nitrogen-based symbioses,phosphorus availability,and accounting for a modern world more productive than the Paleozoic

Evolution of high-productivity angiosperms has been regarded as a driver of Mesozoic ecosystem restructuring. However, terrestrial productivity is limited by availability of rock-derived nutrients such as phosphorus for which permanent increases in weathering would violate mass balance requirements of the long-term carbon cycle. The potential reality of productivity increases sustained since the Mesozoic is supported here with documentation of a dramatic increase in the evolution of nitrogen-fixing or nitrogen-scavenging symbioses, including more than 100 lineages of ectomycorrhizal and lichen-forming fungi and plants with specialized microbial associations. Given this evidence of broadly increased nitrogen availability, we explore via carbon cycle modeling how enhanced phosphorus availability might be sustained without violating mass balance requirements. Volcanism is the dominant carbon input, dictating peaks in weathering outputs up to twice modern values. However, times of weathering rate suppression may be more important for setting system behavior, and the late Paleozoic was the only extended period over which rates are expected to have remained lower than modern. Modeling results are consistent with terrestrial organic matter deposition that accompanied Paleozoic vascular plant evolution having suppressed weathering fluxes by providing an alternative sink of atmospheric CO₂. Suppression would have then been progressively lifted as the crustal reservoir's holding capacity for terrestrial organic matter saturated back toward steady state with deposition of new organic matter balanced by erosion of older organic deposits. Although not an absolute increase, weathering fluxes returning to early Paleozoic conditions would represent a novel regime for the complex land biota that evolved in the interim. Volcanism-based peaks in Mesozoic weathering far surpass the modern rates that sustain a complex diversity of nitrogen-based symbioses; only in the late Paleozoic might these ecologies have been suppressed by significantly lower rates. Thus, angiosperms are posited to be another effect rather than proximal cause of Mesozoic upheaval.     

Origin of dental occlusion in tetrapods: signal for terrestrial vertebrate evolution?

Evolutionary changes of the dentition in tetrapods can be associated with major events in the history of terrestrial vertebrates. Dental occlusion, the process by which teeth from the upper jaw come in contact with those in the lower jaw, appears first in the fossil record in amniotes and their close relatives near the Permo-Carboniferous boundary approximately 300 million years ago. This evolutionary innovation permitted a dramatic increase in the level of oral processing of food in these early tetrapods, and has been generally associated with herbivory. Whereas herbivory in extinct vertebrates is based on circumstantial evidence, dental occlusion provides direct evidence about feeding strategies because jaw movements can be reconstructed from the wear patterns of the teeth. Examination of the evolution of dental occlusion in Paleozoic tetrapods within a phylogenetic framework reveals that this innovation developed independently in several lineages of amniotes, and is represented by a wide range of dental and mandibular morphologies. Dental occlusion also developed within diadectomorphs, the sister taxon of amniotes. The independent, multiple acquisition of this feeding strategy represents an important signal in the evolution of complex terrestrial vertebrate communities, and the first steps in the profound changes in the pattern of trophic interactions in terrestrial ecosystems.     

The origin of herbivory on land： Initial patterns of plant tissue consumption by arthropods

The early fossil record of terrestrial arthropod herbivory consists of two pulses. The first pulse was concentrated during the latest Silurian to Early Devonian （417 to 403 Ma）, and consists of the earliest evidence for consumption of sporangia and stems （and limited fungivore borings）. Herbivorization of most of these tissues was rapid, representing 0 to 20 million-year （m.y.） lags from the earliest occurrences of these organs in the fossil record to their initial consumption （Phase 1）. For approximately the next 75 m.y., there was a second, more histologically varied origination and expansion of roots, leaves, wood and seeds, whose earliest evidence for herbivorization occurred from the Middle-Late Mississippian boundary to the Middle Pennsylvanian （327 to 309 Ma）. The appearance of this second herbivory pulse during the later Paleozoic （Phase 2） is accompanied by major lags of 98 to 54 m.y. between times of appearance of each of the four organ and tissue types and their subsequent herbivory. Both pulses provide a context for three emerging questions. First is an explanation for the contrast between the near instantaneous consumption of plant tissues during Phase 1, versus the exceptionally long lags between the earliest occurrences of plant tissues and their subsequent herbivorization during Phase 2. Second is the identity of arthropod herbivores for both phases. Third is the cause behind the overwhelming targeting of seed-fern plant hosts during Phase 2. Regardless of the answers to these questions, the trace fossil record of plant-arthropod associations provides primary ecological data that remain unaddressed by the body-fossil record alone.     

The earliest evidence of holometabolan insect pupation in conifer wood

Background

The pre-Jurassic record of terrestrial wood borings is poorly resolved, despite body fossil evidence of insect diversification among xylophilic clades starting in the late Paleozoic. Detailed analysis of borings in petrified wood provides direct evidence of wood utilization by invertebrate animals, which typically comprises feeding behaviors.

Methodology/Principal Findings

We describe a U-shaped boring in petrified wood from the Late Triassic Chinle Formation of southern Utah that demonstrates a strong linkage between insect ontogeny and conifer wood resources. Xylokrypta durossi new ichnogenus and ichnospecies is a large excavation in wood that is backfilled with partially digested xylem, creating a secluded chamber. The tracemaker exited the chamber by way of a small vertical shaft. This sequence of behaviors is most consistent with the entrance of a larva followed by pupal quiescence and adult emergence — hallmarks of holometabolous insect ontogeny. Among the known body fossil record of Triassic insects, cupedid beetles (Coleoptera: Archostemata) are deemed the most plausible tracemakers of Xylokrypta, based on their body size and modern xylobiotic lifestyle.

Conclusions/Significance

This oldest record of pupation in fossil wood provides an alternative interpretation to borings once regarded as evidence for Triassic bees. Instead Xylokrypta suggests that early archostematan beetles were leaders in exploiting wood substrates well before modern clades of xylophages arose in the late Mesozoic.     

Carboniferous Onychophora from Montceau‐les‐Mines,France, and onychophoran terrestrialization

The geological age of the onychophoran crown‐group, and when the group came onto land, have been sources of debate. Although stem‐group Onychophora have been identified from as early as the Cambrian, the sparse record of terrestrial taxa from before the Cretaceous is subject to contradictory interpretations. A Late Carboniferous species from the Mazon Creek biota of the USA, Helenodora inopinata, originally interpreted as a crown‐group onychophoran, has recently been allied to early Cambrian stem‐group taxa. Here we describe a fossil species from the Late Carboniferous Montceau‐les‐Mines Lagerstätte, France, informally referred to as an onychophoran for more than 30 years. The onychophoran affinities of Antennipatus montceauensis gen. nov., sp. nov. are indicated by the form of the trunk plicae and the shape and spacing of their papillae, details of antennal annuli, and the presence of putative slime papillae. The poor preservation of several key systematic characters for extant Onychophora, however, prohibits the precise placement of the Carboniferous fossil in the stem or crown of the two extant families, or the onychophoran stem‐group as a whole. Nevertheless, A. montceauensis is the most compelling candidate to date for a terrestrial Paleozoic onychophoran.     

Cord‐forming Palaeozoic fungi in terrestrial assemblages

The fossil record paints a thin picture of early terrestrial life. Useful diagnostic features are rare in the organic‐walled fossils of the first land colonizers, and at first glance the Silurian–Devonian Tortotubus protuberans seems no exception. Now, new material from New York, Gotland and Scotland reveals the ontogenesis and affinity of this problematic organism. Its filamentous early stages (previously referred to Ornatifilum lornensis) demonstrate simple septal perforations and a bilayered cell wall; threads of entwined filaments, bounded by an elaborately sculptured surface, arose via the retrograde growth and subsequent proliferation of secondary branches. This morphology and pattern of growth together indicate an affinity with the ‘higher’ fungi (Dikarya) and document the formation of differentiated mycelium. The presence of complex mycelial fossils in the earliest Silurian corroborates the likely contribution of fungi to the colonization of land and the establishment of modern sedimentological systems; their rise seemingly accompanied the diversification of early embryophytes and the vegetation of the terrestrial biosphere.     

贵州寒武系第二统杷榔组有壳变形虫的新发现*

有壳变形虫(testate amoebae)的演化历史最早可追溯至新元古代早期, 以该时期北美、华北、挪威以及澳洲等多个地区浅海相碳酸盐岩、页岩中发现的瓶状微体化石(vase-shaped microfossils)为标志。此前认为, 显生宙的有壳变形虫最早出现在早泥盆世。长期以来, 早古生代的地层中未发现这类原生生物的明确化石证据。本研究通过对岩石样品进行常规孢粉酸泡分析处理和切磨岩石薄片, 获取原位保存的化石标本的技术方法, 从贵州东部剑河县交榜剖面出露的寒武系杷榔组(第2统第4阶)中获得数枚有壳变形虫(testate amoebae)化石标本。基于标本的显微形态特征, 并结合激光拉曼光谱等研究, 对原先记述为疑源类的Plagasphaera balangensis和P. sp. A两形态种进行重新认识和描述。由于它们在结构和形态上与一些现生的鳞壳虫目(Euglyphida)有壳变形虫极为相似, 因此将先前定为疑源类的Plagasphaera balangensis和P. sp. A两形态属、种名, 分别修订为Palaeoassulina balangensis gen. et sp. nov.和?Palaeoassulina sp. A。该发现不仅将显生宙有壳变形虫的原有化石记录从晚古生代泥盆纪向前延伸至寒武纪早期, 还为调查研究有壳变形虫的系统演化提供关键的生物化石证据。     

Palaeoanellus dimorphus gen. et sp. nov. (Deuteromycotina): a Cretaceous predatory fungus

In habitats where nitrogen is the limiting factor, carnivorous fungi gain an advantage by preying on nematodes and other microorganisms. These fungi are abundant in modern terrestrial ecosystems, but they are not predestined for preservation as fossils. Conclusions on their evolutionary history are therefore mainly based on molecular studies that are generally limited to those taxa that have survived until today. Here we present a fossil dimorphic fungus that was found in Late Albian amber from southwestern France. This fungus possessed unicellular hyphal rings as trapping devices and formed blastospores from which a yeast stage developed. The fossil probably represents an anamorph of an ascomycete and is described as Palaeoanellus dimorphus gen. et sp. nov. Because predatory fungi with regular yeast stages are not known from modern ecosystems, the fungus is assumed to not be related to any Recent carnivorous fungus and to belong to an extinct lineage of carnivorous fungi. The inclusions represent the only record of fossil fungi that developed trapping devices, so far. The fungus lived c. 100 million years ago in a limnetic-terrestrial microhabitat, and it was a part of a highly diverse biocenosis at the forest floor of a Cretaceous coastal amber forest.     

A new genus and species of fossil scorpion (?Euscorpiidae) from the Early–Middle Eocene of Pesciara (Bolca,Italy)

Fossil scorpions are among the oldest terrestrial arthropods known from the fossil record. They have a worldwide distribution and a rich fossil record, especially for the Paleozoic. Fossil scorpions from Mesozoic and Cenozoic deposits are usually rare (except in amber-deposits). Here, we describe the only fossil scorpion from the Early to Middle Eocene Pesciara Lagerstätte in Italy. Eoeuscorpius ceratoi gen. et sp. nov. is probably a genus and species within the family Euscorpiidae. This may be the first fossil record of the Euscorpiidae, which are so far only known from four extant genera. Eoeuscorpius ceratoi gen. et sp. nov. was found in the “Lower Part” of the Pesciara Limestone, which is actually dated Late Ypresian stage (between 49.5 and 49.7 Ma). Besides a possible pseudoscorpion, the here-described fossil scorpion is the second arachnid species known from the Bolca Locality.     

A diverse fossil terrestrial arthropod fauna from New Zealand: evidence from the early Miocene Foulden Maar fossil lagerstätte

Fossil evidence for the evolutionary history of terrestrial arthropods in New Zealand is extremely limited; only six pre‐Quaternary insects (Triassic to Eocene) have been recorded previously, none of Miocene age. The Foulden Maar fossil lagerstätte in Otago has now yielded a diverse arthropod assemblage, including members of the Araneae, Plecoptera, Isoptera, Hemiptera, Coleoptera, Hymenoptera, Trichoptera and Diptera. The fauna significantly emends the fossil record for the Southern Hemisphere, provides an unparalleled insight into a 23‐million‐year‐old New Zealand lake/forest palaeoecosystem and allows a first evaluation of arthropod diversity at a time coeval with or shortly after the maximum marine transgression of Zealandia in the late Oligocene. The well‐preserved arthropods chiefly represent ground‐dwelling taxa of forest floor and leaf litter habitats, mostly from sub‐families and genera that are still present in the modern fauna. They provide precisely dated fossil evidence for the antiquity of some of New Zealand's terrestrial arthropods and the first potential time calibrations for phylogenetic studies. The high arthropod diversity at Foulden Maar, together with a subtropical rainforest flora and fossil evidence for complex arthropod–plant interactions, suggests that terrestrial arthropods persisted during the Oligocene marine transgression of Zealandia.     

The greatest step in vertebrate history: a paleobiological review of the fish-tetrapod transition

Recent discoveries of previously unknown fossil forms have dramatically transformed understanding of many aspects of the fish-tetrapod transition. Newer paleobiological approaches have also contributed to changed views of which animals were involved and when, where, and how the transition occurred. This review summarizes major advances made and reevaluates alternative interpretations of important parts of the evidence. We begin with general issues and concepts, including limitations of the Paleozoic fossil record. We summarize important features of paleoclimates, paleoenvironments, paleobiogeography, and taphonomy. We then review the history of Devonian tetrapods and their closest stem group ancestors within the sarcopterygian fishes. It is now widely accepted that the first tetrapods arose from advanced tetrapodomorph stock (the elpistostegalids) in the Late Devonian, probably in Euramerica. However, truly terrestrial forms did not emerge until much later, in geographically far-flung regions, in the Lower Carboniferous. The complete transition occurred over about 25 million years; definitive emergences onto land took place during the most recent 5 million years. The sequence of character acquisition during the transition can be seen as a five-step process involving: (1) higher osteichthyan (tetrapodomorph) diversification in the Middle Devonian (beginning about 380 million years ago [mya]), (2) the emergence of "prototetrapods" (e.g., Elginerpeton) in the Frasnian stage (about 372 mya), (3) the appearance of aquatic tetrapods (e.g., Acanthostega) sometime in the early to mid-Famennian (about 360 mya), (4) the appearance of "eutetrapods" (e.g., Tulerpeton) at the very end of the Devonian period (about 358 mya), and (5) the first truly terrestrial tetrapods (e.g., Pederpes) in the Lower Carboniferous (about 340 mya). We discuss each of these steps with respect to inferred functional utility of acquired character sets. Dissociated heterochrony is seen as the most likely process for the evolutionarily rapid morphological transformations required. Developmental biological processes, including paedomorphosis, played important roles. We conclude with a discussion of phylogenetic interpretations of the evidence.     

Evolutionary history of regeneration in crinoids (Echinodermata)

The fossil record indicates that crinoids have exhibited remarkable regenerative abilities since their origin in the Ordovician, abilities that they likely inherited from stem-group echinoderms. Regeneration in extant and fossil crinoids is recognized by abrupt differences in the size of abutting plates, aberrant branching patterns, and discontinuities in carbon isotopes. While recovery is common, not all lost body parts can be regenerated; filling plates and overgrowths are evidence of non-regenerative healing. Considering them as a whole, Paleozoic crinoids exhibit the same range of regenerative and non-regenerative healing as Recent crinoids. For example, Paleozoic and extant crinoids show evidence of crown regeneration and stalk regrowth, which can occur only if the entoneural nerve center (chambered organ) remains intact. One group of Paleozoic crinoids, the camerates, may be an exception in that they probably could not regenerate their complex calyx-plating arrangements, including arm facets, but their calyxes could be healed with reparative plates. With that exception, and despite evidence for increases in predation pressure, there is no compelling evidence that crinoids have changed though time in their ability to recover from wounds. Finally, although crinoid appendages may be lost as a consequence of severe abiotic stress and through ontogenetic development, spatiotemporal changes in the intensity and frequency of biotic interactions, especially direct attacks, are the most likely explanation for observed patterns of regeneration and autotomy in crinoids.     

Chemical examinations of some non-vascular Paleozoic plants

The organic chemical constituents of compression fossils ofNematothallus,Spongiophyton, Orestovia andEohostimella are identified and compared with those isolated from living and fossil forms ofBotryococcus (a green alga) andTaeniocrada (a vascular plant fossil). The range and maxima in the carbon numbers observed in the normal, saturated acids isolated fromNematothallus, Orestovia, andSpongiophyton are similar to those of fossilBotryococcus, while those acids contained within compression fossils ofEohostimella are similar to the hydrocarbon composition ofTaeniocrada. Isoprenoid, branched hydrocarbons and steroids identified fromNematothallus, Orestovia, andSpongiophyton suggest these genera have algal affinities, while the presence of thick cuticles and in some cases cutin-like compounds appear to show adaptation to a terrestrial environment. Phenolic compounds retained within rock matrices associated withEohostimella are similar to those isolated fromTaeniocrada suggesting chemical, as well as morphological parallels with the land plant habit. These data are interpreted as indicating an early polyphyletic exploitation of the terrestrial habitat during the Paleozoic.     

Predator–prey interactions amongst Permo‐Triassic terrestrial vertebrates as a deterministic factor influencing faunal collapse and turnover

Unlike modern mammalian communities, terrestrial Paleozoic and Mesozoic vertebrate systems were characterized by carnivore faunas that were as diverse as their herbivore faunas. The comparatively narrow food base available to carnivores in these paleosystems raises the possibility that predator–prey interactions contributed to unstable ecosystems by driving populations to extinction. Here, we develop a model of predator–prey interactions based on diversity, abundance and body size patterns observed in the Permo‐Triassic vertebrate fossil record of the Karoo Basin, South Africa. Our simulations reflect empirical evidence that despite relatively high carnivore: herbivore species ratios, herbivore abundances were sufficient for carnivores to maintain required intake levels through most of the Karoo sequence. However, high mortality rates amongst herbivore populations, even accounting for birth rates of different‐sized species, are predicted for assemblages immediately preceding the end‐Guadalupian and end‐Permian mass extinctions, as well as in the Middle Triassic when archosaurs replaced therapsids as the dominant terrestrial fauna. These results suggest that high rates of herbivore mortality could have played an important role in biodiversity declines leading up to each of these turnover events. Such declines would have made the systems especially vulnerable to subsequent stochastic events and environmental perturbations, culminating in large‐scale extinctions.     

晚三叠世泡孔类苔藓动物Casianopora及其种系发生关系

本文描述了窄唇纲苔藓动物的一个新种Ｃａｓｉａｎｏｐｏｒａｈｅｉｓｐ．ｎｏｖ．,此新种发现于我国滇西北宁蒗彝族自治县松桂组１５层（菊石Ｍａｌａｙｉｔｅｓ－Ｐｓｅｕｄｏｃａｒｄｉｏｃｅｒａｓ带）,是迄今已知的全球三叠纪泡孔类的一个最新代表。重新确定了Ｃａｓｉａｎｏｐｏｒａ属（卡尼中期－诺利早期）的分类位置,它和Ｃｙｓｔｉｔｒｙｐａ属（卡尼早期）都归入Ｆｉｓｔｕｌｉｐｏｒｉｄａｅ科。这两个三叠纪的属,与古生代和古生代后的一些泡孔类的种系发生关系的初步探讨表明,所有古生代后的窄唇纲的各个目（除环口目外）,可能都是从同一演化根系上发展而来的。     

Diversity and significance of late cretaceous permineralized plant remains from Hokkaido,Japan

Middle to Late Cretaceous permineralized plants hitherto described from Hokkaido, Japan are summarized. The fossil flora comprises fungi, ferns, gymnosperms and angiosperms. Many modern fern families have been recognized including Anemiaceae, Cyatheaceae, Dennstaedtiaceae, Gleicheniaceae Loxsomaceae, Lygodiaceae and Matoniaceae. Gymnosperms are most abundant in the flora. Some recently-found materials are tentatively introduced with brief comments emphasizing their morphological and taxonomical significance. A bisporangiate flower ofCycadeoidella japonica Ogura shows fine internal anatomy and provides evidence that the cycadeoidalean ovule was a cupulate, unitegmic structure. Vascular tracheids in the synangial wall support the evolution of cycadeoidalean synangia from Paleozoic seed-fern synangia. A new gymnosperm female fructification has a thick envelope comparable to an angiosperm carpel around a large seed. The angiosperms contain various morphologies that require further extensive study.     

Earliest fossil record of bacterial-cyanobacterial mat consortia: the early Silurian Passage Creek biota (440 Ma, Virginia, USA)

Cyanobacteria in terrestrial and aquatic habitats are frequently associated with heterotrophic bacteria, and such associations are most often metabolically interactive. Functionally, the members of such bacterial–cyanobacterial consortia benefit from diverse metabolic capabilities of their associates, thus exceeding the sum of their parts. Such associations may have been just as ubiquitous in the past, but the fossil record has not produced any direct evidence for such associations to date. In this paper, we document fossil bacteria associated with a macrophytic cyanobacterial mat in the early Silurian (Llandovery) Massanutten Sandstone of Virginia, USA. Both the bacterial and the cyanobacterial cells are preserved by mineral replacement (pyrite subsequently replaced by iron oxyhydroxides) within an amorphous carbonaceous matrix which represents the common exopolysaccharide investment of the cyanobacterial colony. The bacteria are rod-shaped, over 370 nm long and 100 nm in diameter, and occur both as isolated cells and as short filaments. This occurrence represents the oldest fossil evidence for bacterial–cyanobacterial associations, documenting that such consortia were present 440 Ma ago, and revealing the potential for them to be recognized deeper in the fossil record.