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Plant–insect and –fungal interactions in Taxodium-like wood fossils from the Oligocene of southwestern China
Abstract and Figures
Cupressaceae fossil tree stumps from the early Oligocene Lühe coal mine in southwestern China contain abundant quartz-petrified damage traces. The wood fossils were assigned to Taxodioxylon (very similar to extant Taxodium) based on wood anatomy analysis. Within the woods, three types of arthropods- and one fungus-mediated ichnofossils LHIF 1–4 (Lühe wood ichnofossils 1–4) were observed. The boring wood types for LHIF 1–3 are comparable to extant longicorn beetles (Cerambycidae), snout beetles (Curculionidae), and wood wasps (Siricidae). The polyporous structured traces of LHIF 4 were attributed to the invasion of stem canker fungus (Polyporaceae). This first-ever report of Taxodium-like fossil from the Oligocene of southwestern China points to Yunnan serving as a refugium for some lineage of gymnosperms at that time. Furthermore, the extensive traces of arthropods and fungus discovered from the fossil wood have filled a gap in fossil records for insect herbivory in this region. The wood stumps and decomposers suggest a swamp-like environment. The disappearance of Taxodium after the Oligocene supports incremental aridification and changes in winter season temperature conditions, which shifted the late Paleogene mixed deciduous broad-leaved and needle-leaved forest into the present evergreen broad-leaved forest.
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... Mesozoic records include examples in fern and gymnosperm stems from the Triassic (McLoughlin and Strullu-Derrien, 2016) and Jurassic (García et al., 2012;McLoughlin and Bomfleur, 2016;Sagasti et al., 2019;Gee et al., 2022), and in Cretaceous wood and leaves (Brues, 1936;Hower et al., 2013;Donovan et al., 2020;McLoughlin et al., 2021;Philippe et al., 2022). There is also evidence of plant-arthropod-fungal associations in Cenozoic gymnosperm and angiosperm woods and leaves (e.g., Süss and Velitzelos, 2001;McLoughlin, 2020;McLoughlin et al., 2021;Deng et al., 2022). Taken together, these reports form a valuable tool with which to assess the evolutionary history of relationships between plants and certain arthropods and fungi, and to resolve some of the levels of complexity and interconnectedness that existed in past terrestrial ecosystems. ...
... In China, fossil evidence of associations and interactions among plants, arthropods, and fungi comes primarily from the Permian Cathaysia flora and from Oligocene Taxodioxylon Hartig wood (Wan et al., 2016a;Feng et al., 2017;Wei et al., 2019;Deng et al., 2022). Although the Mesozoic plant fossil record from China is extensive (Sze and Li. ...
Dalbergia is a pantropical genus of Fabaceae with a limited fossil record. In this study we describe diverse and well-preserved Dalbergia fossils distributed through Cenozoic (Eocene to Pliocene) strata in Yunnan Province, Southwest China. Comparison with newly collected modern samples from Eurasian forests and herbarium specimens allow accurate identification of our fossil materials, and we demonstrate that Yunnan Province represents an hitherto unrecognised hotspot for the diversification of Dalbergia. Our findings also include evidence of the earliest known mining damage reported on Dalbergia leaflets, which sheds light on their interactions with various arthropods (Leucoptera and other insects) during the early Neogene. As such, our results improve knowledge of Dalbergia fossils in East Asia and are crucial for understanding the evolutionary history of analogous mining behavior observed from modern Dalbergia.
The Paleogene is a crucial period when terrestrial and marine ecosystems recovered from major disruptions and gradually approached their modern states. In the Qinghai-Tibetan Plateau and its surrounding regions, the Paleogene also represents a significant phase of tectonic evolution in the Qinghai-Tibetan Plateau-Himalaya orogeny, reorganization of Asian climates, and evolution of biodiversity. Due to limitations in research conditions and understanding, there are still many controversies regarding stratigraphic divisions in the Qinghai-Tibetan Plateau and its surrounding regions In recent years, extensive studies on sedimentary petrology, magnetostratigraphy, and isotope dating have been conducted in the region. Numerous fossils have been discovered and reported, contributing to a more systematic understanding of biostratigraphy. These studies have laid a solid foundation for the comprehensive investigation of the stratigraphy, biotas and paleogeographic evolution of the Qinghai-Tibetan Plateau and its surrounding regions during the Paleogene. In this paper, we integrate recent research on fossils, isotopic dating, magnetostratigraphy, and geochemistry to refine the stratigraphic divisions and correlation framework of different tectonic units in the region, building upon previous studies. Since the Second Tibetan Plateau Scientific Expedition and Research, the knowledge of Paleogene floras has gradually expanded. This paper discusses the biostratigraphic significance of extinct and newly appeared taxa based on the latest dating results of these plant species. The new understanding of fossil species such as the “Eucalyptus” and Arecaceae establishes connections between the Paleogene flora of the Qinghai-Tibetan region and the biotas of Gondwana, specifically Oceania and South America. The evolutionary history of key taxa near the Yarlung Zangbo suture zone indicates that the collision between the Indian and Eurasian plates occurred approximately 65–54 Ma. Paleoelevation reconstructions, based on plant fossils, suggest that the Hengduan Mountain had already formed their current topographic pattern prior to the Early Oligocene. The warm and humid lowlands adjacent to the main suture zones in the Paleogene Qinghai-Tibetan Plateau served as the primary pathway for biota exchanges. The relatively low elevation of the Himalaya during the Paleogene did not effectively block the moisture from the Indian Ocean.
During the late Eocene to the early Oligocene, marine records document a globally congruent record of declining carbon dioxide concentrations, Antarctic icesheet growth, and associated reorganisation of the global climate system. In contrast, the few existing terrestrial records demonstrate high heterogeneity of environmental change and are difficult to reconcile with those of the oceanic realm. Global drivers for climatic change are particularly difficult to disentangle from regional ones, especially those caused by the complex tectonic evolution of the Tibetan region and its influence on the Asian monsoon system and vegetation. Here, we reconstruct the climatic and environmental history from the late Eocene into the early Oligocene at Lühe Basin, Yunnan, China, a key sedimentary repository along the SE margin of the Tibetan Plateau and an important region for assessing Asian monsoon changes. We investigate a 340-m long section via a multi-proxy approach and climate model simulations. The organic geochemical proxies, via n-alkanes, terpenoids, and hopanes, suggest that thermally immature sediments were deposited in a terrestrial flood plain basin that was primarily occupied by gymnosperms and angiosperms. Branched glycerol diakyl glycerol tetraethers indicate relatively stable temperatures (ca. 10 °C) throughout the section, including across the Eocene-Oligocene boundary. This temperature, cooler than the modern-day average for this site (ca. 15 °C), suggests that this area has not undergone significant uplift since the Oligocene. To further contextualize our data, we tested a suite of climate model simulations with varying pCO2, paleogeography, and Tibetan topography across the Eocene-Oligocene boundary. This data-model comparison suggests that a response to regional factors might explain the absence of a pronounced cooling at Lühe across the Eocene-Oligocene boundary, supporting the emerging picture that the global expression of the EOT in terrestrial environments is more complex than indicated by the marine record.
Mechanisms driving the tectonic evolution of the southeast (SE) margin of Tibet include the Paleogene extrusion of the coherent Indochina lithospheric block and the continuous deformation caused by lower crustal flow since the middle Miocene. The timing and style of regional deformations are keys to determining the role of each mechanism. Fault‐bounded and fault‐controlled Cenozoic basins within the SE margin of Tibet record regional deformation, surface uplift, and variations in paleoclimate but often are poorly dated. New magnetostratigraphy and ⁴⁰Ar/³⁹Ar dating of volcanic ashes constrain precisely the timing of sedimentation within the Lühe Basin to between ~35 and 26.5 Ma. The basin is located in the Chuandian terrane along the Chuxiong fault, which lies ~70 km north of, and parallel to, the Ailao Shan‐Red River fault. The asymmetric syncline of the Lühe Basin suggests syncontractional sedimentation, and the basal age of the basin represents the initiation of the Chuxiong fault and crustal shortening at ~35 Ma. This is coincident with the onset of the Ailao Shan‐Red River fault and supports a kinematic link between them. Our study suggests that, like the Ailao Shan‐Red River fault, the Chuxiong fault is a Paleogene transpressional structure that developed during the extrusion and clockwise rotation of Indochina around the Eastern Himalayan Syntaxis, which caused the late Paleogene deformation and surface uplift of the Chuandian terrane and Indochina. Our revised chronostratigraphy of the Lühe Basin provides further evidence that many of the “Neogene” sedimentary basins in the SE margin of Tibet may be much older than previously thought.
Background:
Chloroplast (cp) genome information would facilitate the development and utilization of Taxodium resources. However, cp genome characteristics of Taxodium were poorly understood.
Results:
We determined the complete cp genome sequences of T. distichum, T. mucronatum, and T. ascendens. The cp genomes are 131,947 bp to 132,613 bp in length, encode 120 genes with the same order, and lack typical inverted repeat (IR) regions. The longest small IR, a 282 bp trnQ-containing IR, were involved in the formation of isomers. Comparative analysis of the 3 cp genomes showed that 91.57% of the indels resulted in the periodic variation of tandem repeat (TR) motifs and 72.46% single nucleotide polymorphisms (SNPs) located closely to TRs, suggesting a relationship between TRs and mutational dynamics. Eleven hypervariable regions were identified as candidates for DNA barcode development. Hypothetical cp open reading frame 1(Ycf1) was the only one gene that has an indel in coding DNA sequence, and the indel is composed of a long TR. When extended to cupressophytes, ycf1 genes have undergone a universal insertion of TRs accompanied by extreme length expansion. Meanwhile, ycf1 also located in rearrangement endpoints of cupressophyte cp genomes. All these characteristics highlight the important role of repeats in the evolution of cp genomes.
Conclusions:
This study added new evidence for the role of repeats in the dynamics mechanism of cp genome mutation and rearrangement. Moreover, the information of TRs and hypervariable regions would provide reliable molecular resources for future research focusing on the infrageneric taxa identification, phylogenetic resolution, population structure and biodiversity for the genus Taxodium and Cupressophytes.
Central Wyoming, USA, was the site of ancient Lake Gosiute during the Early Eocene. Lake Gosiute was a large body of water surrounded by subtropical forest, the lake being part of a lacustrine complex that occupied the Green River Basin. Lake level rises episodically drowned the adjacent forests, causing standing trees and fallen branches to become growth sites for algae and cyanobacteria, which encased submerged wood with thick calcareous stromatolitic coatings. The subsequent regression resulted in a desiccation of the wood, causing volume reduction, radial fractures, and localized decay. The subsequent burial of the wood in silty sediment led to a silicification of the cellular tissue. Later, chalcedony was deposited in larger spaces, as well as in the interstitial areas of the calcareous coatings. The final stage of mineralization was the precipitation of crystalline calcite in spaces that had previously remained unmineralized. The result of this multi-stage mineralization is fossil wood with striking beauty and a complex geologic origin.
We were pleased to see the summary article in Geosciences on fossil wood by George E. [...]
The uplift history of southeastern Tibet is crucial to understanding processes driving the tectonic evolution of the Tibetan Plateau and surrounding areas. Underpinning existing palaeoaltimetric studies has been regional mapping based in large part on biostratigraphy that assumes a Neogene modernisation of the highly diverse, but threatened, Asian biota. Here, with new radiometric dating and newly-collected plant fossil archives, we quantify the surface height of part of Tibet's southeastern margin of Tibet in the latest Eocene (∼34 Ma) to be ∼3 km and rising, possibly attaining its present elevation (3.9 km) in the early Oligocene. We also find that the Eocene-Oligocene transition in southeastern Tibet witnessed leaf size diminution and a floral composition change from sub-tropical/warm temperate to cool temperate, likely reflective of both uplift and secular climate change, and that by the latest Eocene floral modernization on Tibet had already taken place implying modernization was deeply-rooted in the Paleogene.
Under conditions where buried wood is protected from microbial degradation and exposure to oxygen or harsh chemical environments, the tissues may remain unmineralized. If the original organic matter is present in relatively unaltered form, wood is considered to be mummified. Exposure to high temperatures, whether from wild fires or pyroclastic flows, may cause wood to be converted to charcoal. Coalification occurs when plant matter undergoes gradual metamorphosis, producing bituminous alteration products. Examples of all three types of non-mineralized wood are common in the geologic record. This report describes some of the most notable occurrences, reviews past research and introduces data from several localities in North America.
The Ophiostomatales is an Ascomycete order of fungi that accommodates several tree pathogens and many species that degrade wood. These fungi are commonly vectored by Scolytine bark and ambrosia beetles. In recent years it has also been shown that hyperphoretic mites on these beetles can vector some Ophiostomatales. Little is known regarding the Ophiostomatales in China and we have consequently explored the diversity of these fungi associated with conifer-infesting beetles and mites in Yunnan province. Galleries and beetles were collected for 17 beetle species, while 13 mite species were obtained from six of these beetle species. Collectively, 340 fungal isolates were obtained, 45 from beetles, 184 from mites, 56 from galleries and 55 isolates where the specific niche was not clear. DNA sequences for five gene regions (ITS, LSU, BT, EF, and CAL) were determined for fungal isolates representing different morphological groups. Phylogenetic analyses confirmed the presence of 19 fungal taxa, including five novel species described here as Ophiostoma acarorum sp. nov., Ophiostoma brevipilosi sp. nov., Graphilbum kesiyae sp. nov., Graphilbum puerense sp. nov., and Leptographium ningerense sp. nov. Ophiostoma ips was the most frequently isolated species, representing approximately 31% of all isolates. Six of 19 taxa were present on mites, beetles and in the galleries of the beetles, while three species were found on mites and galleries. Two species were found only on mites and one species only on a beetle. Although the numbers of beetles and mites were insufficient to provide statistical inferences, this study confirmed that mites are important vectors of the Ophiostomatales in China. We hypothesize that these mites are most likely responsible for horizontal transfer of fungal species between galleries of different beetle species. The fact that half of the fungal species found were new to science, suggests that the forests of east Asia include many undescribed Ophiostomatales yet to be discovered.
Palaeoendemic plants, whose past geographical distribution was much larger than their present distribution ranges, play an important role in reconstructing biogeographical events, and in tracking floristic relationships among continents throughout geological time. Improving knowledge about their geographical histories helps to understand the evolution of biodiversity under palaeoenvironmental changes. Here, we report the first fossil record of Dipteronia from East Asia, namely Dipteronia brownii McClain et Manchester, based on diagnostic winged fruit remains from the Rupelian (c. 32 ± 1 Ma) of the Lühe Basin in southwestern China. The fossil species is characterized by (i) a samaroid mericarp, with an orbicular wing, (ii) a primary vein extending from the attachment scar, and ramifying into secondary and tertiary veins that radiate out from the seed body area and dichot-omize toward the mericarp. Dipteronia was widely distributed in the Northern Hemisphere during the Palaeogene and is now endemic to central and southwestern China with two extant species. This new finding significantly expands the historical distribution of Dipteronia into China by the early Oligocene, and its present-day endemism is apparently the result of extirpation from North America and the higher latitudes of Asia associated with Cenozoic climatic changes.
For the first time, a synthesis on the research work done in Europe on all Bark And Wood Boring Insects In Living Trees (BAWBILT) is presented. As final product of a four-year research project gathering together 100 scientists from 24 countries, the book is the fruit of a real collective synthesis in which all European specialists have participated. It reviews and comments on all the European literature, while considering the biological (trees, insects, associated organisms, and their relationships) and forest management aspects. However, although focused on the European forest, it also compares the available information and interpretations to those concerning similar species in other continents. It ends with propositions of research priorities for Europe. The book is directed to all scientists and students concerned with forest entomology and ecology, as well as to forest managers and all scientific public interested in forest biology.
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.
Two New Caledonian conifers, Dacrydium guillauminii and Retrophyllum minor, both of Podocarpaceae, occupy distinctive habitats in which roots are characteristically inundated. SEM studies of stem wood anatomy of both reveal characteristic vertical intercellular channels, not apparent in light microscope examination. These channels are regular in spacing as seen in radial sections and indefinite in vertical extent; they were observed only adjacent to rays. These channels, not reported in other conifers, are a potential source of aeration for roots and submersed stem portions. Other wood details are reported. In order to provide a more comprehensive picture of adaptation to these habitats, observations based on field work are provided: these conifers are not only adapted to little-varying inundation of stems and lower roots, they are subjected to much greater sunlight than are upland conifers of New Caledonia.
Fossil Sequoia in Yunnan Province, China during the Miocene is here reported for the first time. The morphological features of shoots and cuticle are described. Its historical and geological distribution is also discussed. According to the characteristics of the assemblage of spore and pollen in the fossil collection site and the nature environment of the native S. sempervirens lived in American, the discovery of the fossil Sequoia indicated the presence of moist subtropical climate in Yunnan Province during the Miocene.
full journal issue presenting new list of agreed upon features for taxonomic work
Inside Wood is an Internet-accessible wood anatomy reference, research, and teaching tool. The Inside Wood database has coded wood anatomical descriptions based on the IAWA List of Microscopic Features for Hardwood Identification and is accompanied by a collection of photomicrographs. As of November 2010 there were over 5,800 descriptions and 36,000 images of modern woods, and over 1,600 descriptions and 2,000 images of fossil woods. CITES-listed timber species and other endangered woody plants are included in this digital collection hosted by North Carolina State University's library. This web site has value in helping with wood identification because it has a multiple entry key that allows searching by presence or absence of IAWA features and it serves as a virtual reference collection whereby descriptions and images can be retrieved by searching by scientific or common name or other keywords.
I. II. III. IV. V. References SUMMARY: We review the literature on host resistance of ash to emerald ash borer (EAB, Agrilus planipennis), an invasive species that causes widespread mortality of ash. Manchurian ash (Fraxinus mandshurica), which coevolved with EAB, is more resistant than evolutionarily naïve North American and European congeners. Manchurian ash was less preferred for adult feeding and oviposition than susceptible hosts, more resistant to larval feeding, had higher constitutive concentrations of bark lignans, coumarins, proline, tyramine and defensive proteins, and was characterized by faster oxidation of phenolics. Consistent with EAB being a secondary colonizer of coevolved hosts, drought stress decreased the resistance of Manchurian ash, but had no effect on constitutive bark phenolics, suggesting that they do not contribute to increased susceptibility in response to drought stress. The induced resistance of North American species to EAB in response to the exogenous application of methyl jasmonate was associated with increased bark concentrations of verbascoside, lignin and/or trypsin inhibitors, which decreased larval survival and/or growth in bioassays. This finding suggests that these inherently susceptible species possess latent defenses that are not induced naturally by larval colonization, perhaps because they fail to recognize larval cues or respond quickly enough. Finally, we propose future research directions that would address some critical knowledge gaps.
© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.
Summary
Supplemented by information from other organs, such as cones, shoots, leaves and dispersed pollen, anatomical analysis of autochthonous coalified woods and research into dispersed cuticles provides data for reconstructing Miocene peat-swamp vegetations in Central Europe. The present paper is focussed on the description and taphonomical interpretation of conifer remains from the Middle Miocene Second Seam horizon of the Lusatian brown coal field in the Lower Lusatia region (Brandenburg, Germany) exposed in the ancient opencast mine “Meuro” north of Senftenberg.
Wood-anatomical evidence of occurrence of the genus Sciadopitys is established by the presence of Sciadopityoxylon wettsteinii JURASKY. In addition, remains published as Sciadopitys tertiaria MENZEL (leaves, cuticles and male cones), Sciadopityspollenites RAATZ and a part of rootlets summarized in the morphotaxon Marcoduria inopinata WEYLAND can be assigned to this genus. The taphonomical dispersion of Sciadopitys remains in the sequence root – trunk – branch – cone – leaf – pollen corresponds to the petrographical turn humotelinitic coal (root coal = “flaserige Kohle” – xylitic coal – “needle” coal) – humodetrinitic coal. The bases of the slender, up to 2.6 m high and relatively small in diameter, tree trunks are sunken into the substratum, which was water-saturated and soft during burial.
The presence of the genus Cunninghamia could be established after a reconsideration of the botanical affinity of wood remains conventionally classified as Glyptostroboxylon tenerum (KRAUS) CONWENTZ. Supporting taxonomic evidence is provided by the occurrence of a short-needled twig of Cunninghamia miocenica ETTINGSHAUSEN showing histologic characteristics. Cuticles frequently occur whereas Cunninghamiaepollenites NAGY was identified only sporadic. Taphonomically, the prevalence of thoroughly dispersed and decomposed cuticle remains in light coloured humodetrinitic, fusite-bearing brown coal is characteristic of stagnating peat accumulation.
In addition to wood remains identified as Taxodioxylon taxodii GOTHAN, the genus Taxodium is represented by the fossil species Taxodium dubium (STERNBERG) HEER (twigs, cones, seeds), as well as Enormicutis angusticavata SCHNEIDER (dispersed cuticles). Pollen belonging to Taxodium is included in the morphotaxa Inaperturopollenites concedipites (WODEHOUSE) KRUTZSCH und I. hiatus (R. POTONIÉ) THOMSON & PFLUG. The taphonomy of the genus is particularly characterized by the preservation of massive stumps around and above a sand bar, like intercalation in the brown coal, cuticulae dispersae in the level of clastic sediments, and peaks of Inaperturopollenites. A Nyssa-Taxodium-association is proved in the underlying bed. In contrast to the Glyptostrobus-swamp, only the stumps may be regarded as contribution for peat formation.
Wood-anatomical identification of five distinctive Pinuxylon species provides evidence of five species of the genus Pinus. An about corresponding number of well characterized morphospecies of dispersed cuticles includes a new species: Pinicutis bergheideana SCHNEIDER. The Pinus-facies defined for the Second Lusatian seam horizon is characterized by three-needled short shoots (Pinus cf. rigios [UNGER] ETTINGSHAUSEN sensu MAI), Pinicutis hexacytica SCHNEIDER, cone axes from Pinus spinosa HERBST and/or P. ungeri STUR, “Buchtenkork”/indented cork and pollen of Pitysporites labdacus (R. POTONIÉ) THOMSON & PFLUG. The Xylotomical taxa Pinuxylon parryoides (GOTHAN) KRAEUSEL emend. VAN DER BURGH and P. ponderosoides VAN DER BURGH are components of this taphocoenosis. The taxonomical relations need further investigations. In the petrographical turn root coals are not developed. The lack of Pinus-stumps in the Second Lusatian seam horizon is explained by the permanence in the area of the primary oxidation zone (acrotelm).
Comparison with the wood of the extant species Cathaya argyrophylla indicates that fossil material from the brown coal mine “Meuro” conventionally identified as Piceoxylon pseudotsugae GOTHAN emend. VAN DER BURGH represents the genus Cathaya. The wood co-occurs with the morphotaxa Cathaya roseltii SCHNEIDER (needles) and Cathayapollis ZIEMBIŃSKA-TWORZYDŁO (pollen). Taphonomical characteristics and petrography of Cathaya–remains display analogies to those of Pinus. A taphocoenosis in which the Cathaya-morphotaxa plays a distinguishing role is recognized.
Wood remains described in the present paper enlarge the stock of characteristics for facies types (especially A, P, M) defined by SCHNEIDER (1969a, b) for the Second Lusatian seam. The following characteristics are correlated with the botanical affinity:
• morphology (stump, trunk, branch),
• dimension (height, diameter),
• petrographical variety (fibrous xylite, “Bruchxylit”, gelified xylite),
• petrography (colour, structure) of surrounding coal matrix.
The xylotomical taxa are assigned to a facies type and a phytocoenosis on the basis of the results of taphonomic analysis. The ombrotrophical conditions increasing in the course of K-A-P-M succession are indicated by increasing in height/diameter quotient of the stumps as function of ascending water saturation resp. reducing stability of the peat substratum.
Key words: Autochthonous fossil woods – cuticle investigations – palynology - succession stages – taphonomy – Miocene - Lower Lusatia (Brandenburg).
Fourteen oribatid mite fossils have been recovered from a terrestrial Devonian deposit near Gilboa, New York; they allow an approximate doubling of the known age of the group. Two species are represented by specimens complete enough to allow descriptions. Protochthonius gilboa n.gen. and sp. is the earliest derivative known member of the Enarthronota, and comprises the monobasic new family Protochthoniidae. Devonacarus sellnicki n.gen. and sp. also represents a monobasic new family, Devonacaridae; it may also be an early derivative enarthronote mite, but its relationships are uncertain. Two other species are represented at the site, but meaningful descriptions are not possible with available material. -Authors
Latest Middle Pennsylvanian, lowland, equatorial communities of Euramerica occurred in equable, wetland habitats with peat substrates dominated by pole-like lycopod trees amid subdominant tree ferns and seed ferns, along with sphenopsids and rare cordaites. In these taxonomically diverse and physiognomically varied communities detritivores overwhelmingly channeled plant biomass from primary production to reduced organic matter. However, by the Late Pennsylvanian, when Psaronius tree ferns replaced lycopods as dominant floral elements, there was a dramatic increase in the spectrum and intensity of herbivore functional-feeding-groups. This shift toward the importance of herbivores is supported by a significant increase of arthropod coprolites in peat litter whose origin is most parsimoniously explained by herbivory. Equally important are the earliest documented occurrences of highly stereotyped and host specific plant damage caused by herbivorous insects. We describe two of these distinctive interactions as the newly established ichnogenera Pteridiscaphichnos, a detritivore-created, tunnel-and-gallery network in Middle- and Late Pennsylvanian Psaronius stems; and Pteriditorichnos, an elongate petiole gall from the frond of the Late Pennsylvanian Psaronius chasei, caused by an herbivorous and endophytic larval holometabolan. The Psaronius stem-boring, Pteridiscaphichnos, occurs in the late Middle Pennsylvanian form "layered cells morphotype" and the Late Pennsylvanian species Psaronius magnificus and P. chasei. This distinctive stem boring is characterized by evacuated ground parenchyma replaced by frass of ellipsoidal coprolites 1.0 mm x 3.0 mm in average size and macerated fragments of processed but unconsumed parenchyma. All known specimens occur among linear to arcuate vascular strands interior to the outer sclerenchyma cylinder. The lack of reaction tissue indicates postmortem consumption of relatively soft, nutrient-rich parenchyma, although construction of the tunnel-and-gallery network preceded significant decomposition of the trunk since lignified tissues, such as sclerenchyma, vascular tissue and part of the root mantle, were still structurally sound and were avoided by the detritivores. Based on the geometry of the boring, coprolite structure, and the life-habits of modern ecologic analogs inhabiting tree fern trunks, the culprit of Pteridiscaphichnos is attributed to a roach or roach-like insect. The Late Pennsylvanian gall, Pteriditorichnos, is known only on Psaronius chasei hosts, and occurs in basal frond petioles. The structure of this gall consists of a lumen formed by removal of ground parenchyma through the petiolar axis, and its partial replacement by files of hypertrophied cells organized into radiating, impinging tufts that define a differentiated zone of callus. The lumen, encapsulated by callus, contains regions with frass consisting of coprolites and macerated parenchyma fragments. Barrel-shaped coprolites average 1.5 mm x 2.5 mm in size, have smooth longitudinal surfaces but ragged ends, and contain recognizable cell types found in the surrounding callus and unmodified parenchyma. All lines of evidence indicate that an endophytic insect tunneled through medullary petiolar parenchyma, which induced plant response by production of callus, and was followed by insect consumption of the callus. Based on the three-dimensional structure of this gall, the particulate nature of the coprolites, and an extensive literature documenting known gall-making arthropods, we identify the culprit as a holometabolous larva. Although this earliest known gall resembles those made by extant tenthredinoid Hymenoptera, it is unassignable to any modern lineage of Holometabola. Although possible body fossils of the Holometabola have been mentioned in Pennsylvanian strata, Pteriditorichnos provides the first high-resolution, trace-fossil documentation of the life-habits of any Paleozoic larval holometabolan. Because of its exceptional anatomical preservation and Late Pennsylvanian age, this larval-induced gall is important in assessing existing hypotheses regarding the ancestral life-habits of the earliest larval Holometabola. Evidence indicates: 1. That the larval holometabolan galler occurred in a climatically equable, tropical, and humid swamp forest with a peat substrate. 2. That endophytic petiole galling is the earliest known life-habit for the Holometabola, and that stem galling or a related endophytic guild was an ancestral/behavior for this clade. 3. The prediction of LAMEERE, later elaborated by MALYSHEV, is vindicated: the endophytic penetration of tissues in primitive vascular plants (especially pteridophytes) was integrally associated with the holometabolous condition. These data indicate not only the occurrence of a stem borer and a herbivore galler during the Paleozoic, but also document the presence of an intimate interaction between a herbivore and its responding plant host as early as the Late Pennsylvanian.
Several studies provided evidence that family-level insect diversity remained flat throughout the initial mid-Cretaceous angiosperm radiation 125–90 million years ago. As this result has engendered considerable commentary, a reanalysis was done of a new dataset of 280 plant-associated insect families spanning the 174 million year interval of the Jurassic–Paleogene periods from 201 to 23 million years ago. Lineage geochronologic ranges were determined, and feeding attributes were characterized by: (i) dominant feeding guild (herbivore, pollinator, herbivore–pollinator, pollinator–mimic, xylophage); (ii) membership in one of eight functional feeding groups; and (iii) dominant plant host or host transition (cryptogam/fern only, cryptogam/fern → angiosperm, gymnosperm only, gymnosperm → angiosperm, angiosperm only). A time-series plot of insect lineages and their dominant plant–host affiliations resulted in four conclusions. First, insect lineages with dominant gymnosperm hosts reached a level of 95 families in the 35 million years preceding the initial angiosperm radiation. Second, earlier insect lineages with gymnosperm → angiosperm host transitions and newly originated insect lineages that developed dominant associations with emerging angiosperms rapidly diversified during the angiosperm radiation, later establishing a plateau of 110 families during a 20 million year interval after the initial angiosperm radiation. Third, these two diversity maxima were separated during the angiosperm radiation by a diversity minimum, the Aptian–Albian gap, indicating major turnover and time-lag effects associated with the extirpation and acquisition of plant associations. Last, insect lineages most affected during this interval were herbivores and pollinators, exophagous feeders, and those hosting gymnosperms, angiosperms and gymnosperm → angiosperm transitions. These data largely explain the flat or even decreased level of insect diversity immediately before, during, and after the initial angiosperm radiation.
Although oribatid mites are essential to the decomposition of plant tissues in modern temperate forests by assisting conversion of primary productivity to soil organic matter, little is known of their paleoecologic history. Previously there has been scattered and anecdotal evidence documenting oribatid mite detritivory in Pennsylvanian plant tissues. This study evaluates the incidence of oribatid mite damage for seven major coal-ball deposits from the Illinois and Appalachian sedimentary basins, representing a 17 million year interval from the Euramerican tropics. Although this interval contains the best anatomically preserved plant tissues with oribatid mite borings in the fossil record, coeval oribatid mite body-fossils are absent. By contrast, the known body-fossil record of oribatid mites commences during the Middle Devonian, but does not reappear until the Early Jurassic, at which time mite taxa are modern in aspect. All major plant taxa occurring in Pennsylvanian coal swamps, including lycopsids, sphenopsids, ferns, seed ferns and cordaites, were consumed by oribatid mites. Virtually every type of plant tissue was used by mites, notably indurated tissues such as bark, fibrovascular bundles and especially wood, as well as softer seed megagametophytic and parenchymatic tissues within stems, roots and leaves. Significant evidence also exists for secondary consumption by mites of tissues in macroarthropod coprolites. Our data indicate that oribatid mites consumed dead, aerially-derived plant tissues at ground level, as well as root-penetrated tissues substantially within the peat. Oribatid mites were important arthropod decomposers in Pennsylvanian coal swamps of Euramerica. The wood boring functional-feeding-guild was expanded by insects into above-ground, live trees during the early Mesozoic. New food resources for insect borers resulted from penetration of live tissues such as cambium and phloem, and the invasion of heartwood and other hard tissues mediated by insect-fungus symbioses. Termites and holometabolous insects were prominent contributors to this second wave of wood-boring, exploiting gymnosperms and angiosperms as both detritivores and herbivores. An earlier emplacement of oribatid mites as detritivores of dead plant tissues continued to the present, but without a documented trace-fossil record.
Premise of the study: The paleogeographical origin of the relict North American Sequoia sempervirens is controversial. Fossil records indicate a Neogene origin for its foliage characteristics. Although several fossils from the Miocene sediments in eastern Asia have been considered to have close affi nities with the modern S. sempervirens , they lack the typical features of a leafy twig bearing linear as well as scale leaves, and the fertile shoots terminating by a cone. The taxonomic status of these fossils has
remained unclear.
• Methods: New better-preserved fossils from the upper Miocene of China indicate a new species of Sequoia . This fi nding not only confi rms the former presence of this genus in eastern Asia, but it also confi rms the affi nity of this Asian form to the modern relict S. sempervirens .
• Key results: The principal foliage characteristics of S. sempervirens had already originated by the late Miocene. The eastern Asian records probably imply a Beringian biogeographic track of the ancestor of S. sempervirens in the early Neogene, at a time when the land bridge was not too cool for this thermophilic conifer to spread between Asia and North America.
• Conclusions: The climatic context of the new fossil Sequoia in Southeast Yunnan, based on other fl oristic elements of the fossil assemblage in which it is found, is presumed to be warm and humid. Following the uplift of the Qinghai-Tibet Plateau, this warm, humid climate was replaced by the present monsoonal climate with dry winter and spring. This change may have led to the disappearance of this hygrophilous conifer from eastern Asia.
Archips goyerana, new species, is described and illustrated from southern Louisiana and southwestern Mississippi. It appears to be host-specific on baldcypress (Taxodium distichum; Taxodiaceae) and has become a serious pest of that tree species since its discovery in 1983. Notes on its biology are given, and differences between it and its sister species, A. argyrospila, are outlined. Although morphological differences between A. goyerana and A. argyrospila are subtle, they are corroborated by differences in forewing pattern, larval food plant preferences, and molecular data.
Results of a 4 year survey on the larval and pupal stages of Aenetus virescens are presented and supplemented by observations on the duration and seasonal abundance of the egg, larva, pupa, and adult. The length of the life cycle varied from about 1–4 years. Adults emerge mainly in spring and early summer (September–December). Eggs mature within 1 month and young larvae feed on dead wood and fungi for about 2 months. New tunnels were constructed in trees throughout the year but the numbers peaked in January–March. New pupae were observed from March–November. The pupal stages that began after June lasted for a progressively shorter time.
Two new wood soldier flies, Clemoxyla aculeolata gen. et sp. nov. and Archeoxyla gigantea gen. et sp. nov., are described from mid-Cretaceous Myanmar (Burmese) amber. Clemoxyla gen. nov. is most characteristic of the broadened cell d and presence of short M1+2 stem after the cell d. The apical palpus of Archeoxyla gen. nov. is obviously expanded that is distinct from other known fossil xylomyid genera. Based on the analysis of key features of body and venation, the genus Cretarthropeina Solórzano Kraemer and Cumming, 2019 is transferred from Xylomyidae to Stratiomyidae as the sixth soldier fly from mid-Cretaceous Myanmar amber. In addition, a key to the genera of all known fossil and extant xylomyids is provided.
Longitudinally aligned borings attributed to the ichnotaxon Dekosichnus meniscatus in the inner secondary wood of a silicified Middle–Late Jurassic conifer from Argentina contain finely granular frass particles arranged in meniscoid laminae. Synchrotron X-ray computed tomographic reconstruction of the borings reveals new characters of this ichnotaxon, such as opposing orientations of menisci in some adjacent borings, regular spacing of minor and major meniscoid laminae, a scarcity of tunnel branching, and rare occurrences of cylindrical–spherical terminal chambers on excavations. Architectural and distributional features of the galleries suggest excavation by cerambycid beetle larvae, thus representing one of the earliest potential fossil records of this group. The borings are confined to the inner wood of a young tree that experienced a moderately seasonal climate in a volcanically influenced landscape. By detecting subtle heterogeneities in composition, this study demonstrates that high-energy synchrotron X-ray tomography can characterize anatomical features and complex ecological interactions within even densely permineralized (silicified) plant fossils.
Permineralized woods of Nothofagoxylon scalariforme from Miocene strata (Río Leona Formation) that crop out in the southwestern region of Santa Cruz Province, Argentina were studied. Six types of borings are described, filled with coprolites or homogeneous excrement masses (frass). Based on size, shape, orientation and content, the borings can be related to oribatid mites (Acari: Oribatida), beetles (Insecta: Coleoptera) and springtails (Insecta: Collembola). The host woods show fungal decay and are colonized by hyphae and spores, which suggest various forms of interaction with the arthropods and the Nothofagoxylon trees. The possible types of interactions developed between different organisms involved in the excavation, consumption and degradation of fossil woods in Miocene forests of Patagonia are discussed. Collectively, this evidence indicates that the analyzed woods were in an advanced state of decay prior to their fossilization and that several groups of organisms were active participants in the decomposition of woody substrates, or used them as nesting places, in early Neogene forest ecosystems of Patagonia.
The early Oligocene represents the beginning of the modern icehouse world. To better understand how vegetation and climate changed during this period, we reconstruct quantitively the early Oligocene vegetation and climate by analyzing fossil spore and pollen assemblages from Lühe basin, Yunnan Province, in southwestern China. The reconstructed paleoclimate using Bioclimatic Analysis shows that the mean annual temperature (MAT) ranged from 14.4 to 15.4 °C, the mean annual precipitation (MAP) was 1070.1–1297.8 mm, and the range of mean temperature of coldest quarter (MTCQ) was 6.7 to 7 °C, lower than that of the present day (9.3 °C). The reconstructed regional vegetation indicates a deciduous broadleaved forest mixed with evergreen broadleaved elements. Considering other Paleogene floras in Yunnan, deciduous broadleaved elements dominated in this area from the late Eocene to the early Oligocene. The paleoclimate reconstruction suggests that this vegetation composition may have been controlled mainly by cold winter temperatures.
Herbivore damage patterns on fossil leaves are essential to explore the evolution of plant-herbivore interactions under paleoenvironmental changes and to better understand the evolutionary history of terrestrial ecosystems. The Eocene–Oligocene transition (EOT) is a period of dramatic paleoclimate changes that significantly impacted global ecosystems; however, the influences on plant-herbivore interactions during this period are largely unknown. We identified taxonomic composition of the flora, and investigated well-preserved herbivore damage on fossil leaves from two layers of the Lawula Formation in Markam County, southeastern Qinghai-Tibetan Plateau (QTP), China. Besides, paleoclimate conditions were reconstructed using fossil plant assemblages. The plant assemblage from the latest Eocene layer (MK-3, ~34.6 Ma) was dominated by Fagaceae and Betulaceae, whereas Rosaceae and Salicaceae were the most abundant in the earliest Oligocene layer (MK-1, ~33.4 Ma). In MK-3, 932 out of 2428 fossil leaves were damaged and presented 41 damage types (DTs). The richest functional feeding groups (FFGs) in this layer were hole feeding, margin feeding, and galling. In MK-1, 144 out of 599 leaves were damaged and presented 20 DTs, with the major FFGs being hole feeding, margin feeding, and skeletonization. Generally, MK-3 had a significantly higher damage frequency (DF) and more DTs compared to MK-1. The decline in temperature, accompanied by the mountain uplift during the EOT on the QTP margin, led to changes in plant composition, with a consequent decrease in herbivory quantity and diversity. Our results shed new light on the influence of paleoenvironmental changes in shaping the evolution of biodiversity as well as the ecosystem on the plateau.
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).
The southeastern margin of the Tibetan Plateau (TP) is a key region for understanding the region's surface uplift mechanisms. This study focused on the Jiuziyan and Shuanghe Formations (Fms) in the Jianchuan Basin, both of which include lacustrine calcareous mudstones and marls. Ostracods of the genus Austrocypris found within the Jiuziyan and Shuanghe Fms constrain the age of strata to the Late Eocene. This study used two different proxies, i.e. fossil pollen coexistence and the δ^(18)O (VPDB) values of carbonate (δ^(18)O_c), to reconstruct paleoelevation and the extant paleoenvironment from lacustrine calcareous mudstones and marls preserved in the Eocene stratigraphy of the Jianchuan Basin. The coexistence approach (CA) using pollen data from the Shuanghe Fm indicates a paleoelevation of 1.3–2.6 km above sea level (asl), which would most probably have been associated with a vegetation cover consisting of tropical-subtropical, deciduous, coniferous, broadleaf forests. The reconstructed mean annual air temperature (MAAT) had a value of 16.8–21.7 °C, warmer than today's MAAT (~6 °C). Oxygen isotope results from the Jiuziyan Fm, with/without modification between Eocene and modern Myanmar sea level, suggested that the surface of the Jianchuan Basin was at a paleoelevation between 0.5^(+0.8)_(–0.5) km asl and 2.5 ± 0.7 km asl (δ^(18)Omw: −8.9 ± 1.3‰, 2σ). During the Shuanghe Fm sedimentation the paleoelevation was between 0.9^(+0.7)_(–0.7) km asl and 2.9 ± 0.6 km asl (δ^(18)O_(mw): −9.5 ± 1.1‰, 2σ). Our results suggest that a stepwise uplift of Jianchuan Basin and crustal thickening initiated during the Eocene was the cause of passive surface uplift of the southeastern Tibetan Plateau (TP) rather than Miocene lower crustal flow.
Yunnan, in southwestern China, straddles two of the world's most important biodiversity hot spots (i.e., a biogeographic region that is both a reservoir of biodiversity and threatened with destruction) and hosts more than 200 fossiliferous sedimentary basins documenting the evolutionary history of that biodiversity, monsoon development, and regional elevation changes. The fossil biotas appear modern and have been assumed to be mostly Miocene in age. Dating has been by cross-correlation using palynology, magnetostratigraphy, and lithostratigraphy because numerical radiometric ages are lacking. Here we report the first unequivocal early Oligocene age (33-32 Ma) of a section in the Lühe Basin (25.141627°N, 101.373840°E, 1890 m above mean sea level), central Yunnan, based on U-Pb zircon dates of unreworked volcanic ash layers in a predominantly lacustrine succession hosting abundant plant and animal fossils. This section, located in Lühe town, is correlated with an adjacent section in the Lühe coal mine previously assigned to the upper Miocene based on regional lithostratigraphic comparison. Our substantially older age for the Lühe town section calls into question previous estimates for the surface uplift and climate history of the area, and the age of all other correlative basins. The modernization of the biota ~20 m.y. earlier than previously thought overturns existing concepts of vegetation history in southwestern China, and points to Paleogene modernization of the biota in Yunnan and associated Asian biodiversity hot spots.
Fossils document the existence of trees and wood-associated organisms from almost 400 million years ago, and today there are between 400,000 and 1 million wood-inhabiting species in the world. This is the first book to synthesise the natural history and conservation needs of wood-inhabiting organisms. Presenting a thorough introduction to biodiversity in decaying wood, the book studies the rich diversity of fungi, insects and vertebrates that depend upon dead wood. It describes the functional diversity of these organisms and their specific habitat requirements in terms of host trees, decay phases, tree dimensions, microhabitats and the surrounding environment. Recognising the threats posed by timber extraction and forest management, the authors also present management options for protecting and maintaining the diversity of these species in forests as well as in agricultural landscapes and urban parks.
Late Miocene woods were investigated from the Lühe Basin in Chuxiong Borough, central Yunnan, China. The calcified woods preserved in the Shigucun member of the Shihuiba Formation, are represented by fallen logs and stumps. Two species of fossil wood, Taxodioxylon cryptomeripsoides Schonfeld 1953 and T. cunninghamioides Watari 1948, are described. Their anatomical structure shows striking similarities to the woods of Cryptomeria and Cunninghamia respectively. That the two fossil woods are classed in the Taxodiaceae suggest a subtropical, humid, and warm environment in Lühe during Late Miocene. They compare favorably to other fossil specimens and species reported from localities ranging from Cretaceous to Pliocene. This is the first record of the presence of the species T. cryptomeripsoides and T. cunninghamioides in Late Miocene of Yunnan.
Cenozoic plant relicts are those groups that were once widespread in the Northern Hemisphere but are now restricted to some small isolated areas as a result of drastic climatic changes. They are good proxies to study how plants respond to climatic changes since their modern climatic requirements are known. Herein we look at the modern distribution of 65 palaeoendemic genera in China and compare it with the Chinese climatic pattern, in order to find a link between the plant distribution and climate. Central China and Taiwan Island are shown to be diversity centres of Cenozoic relict genera, consistent with the fact that these two regions have a shorter dry season with comparatively humid autumn and spring in China. Species distribution models indicate that the precipitation parameters are the most important variables to explain the distribution of relict genera. The Cenozoic wide-scale distribution of relict plants in the Northern Hemisphere is therefore considered to be linked to the widespread humid climate at that time, and the subsequent contraction of their distributional ranges was probably caused by the drying trend along with global cooling.
New observations on burrowing behavior of mayfly larvae are given for the leptophlebiids Paraleptophlebia packi and P. bicornuta in North America, and Jappa kutera in Australia; for the potamanthids Potamanthus idiocerus in Taiwan, and P. formosus and Rhoennanthus speciosus in Malaysia; for the polymitarcyids Proboscidoplocia spp. in Madagascar, Afroplocia sampsoni in South Africa, and Ephoron album in North America; and for the ephemerids Ephemera simulans and Litobrancha recurvata in North America, and Palingenia fuliginosa in east Europe. Paraleptophlebia packi forms burrows in silt, whereas P. bicornuta is an interstitial dweller. Potamanthus idiocerus and R. speciosus are the first species of Potamanthidae known to form burrows in silt; however, P. formosus is more typical of the family in that it is an interstitial dweller. Silt burrows made by Leptophlebiidae and Potamanthidae are formed along a rock interface and are never U-shaped, but those formed by advanced burrowers in the Polymitarcyidae and Ephemeridae are independent of rocks and often U-shaped. New evidence of burrowing in plesiotypic polymitarcyid lineages with flat-bodied larvae, represented by Proboscidoplocia and Afroplocia, is provided. Ephgoron album is a highly flexible burrower; its larvae form burrows in depositional substrates, but are interstitial dwellers in erosional substrates. Palingenia fuliginosa is the first non-polymitarcyid burrower to be found burrowing in wood.
Climate change during the Quaternary played an important role in the distribution of extant plants. Herein, cone scales of Cedrus (Pinaceae) were uncovered from the Upper Pliocene Sanying Formation, Longmen Village, Yongping County of Yunnan Province in southwestern China. Detailed comparisons show that these fossils all belong to the genus Cedrus (Pinaceae), and a new species is proposed, Cedrus angusta sp. nov. This find expands the known distribution of Cedrus during the Late Pliocene to Yunnan, where the genus no longer exists in natural forests. Based on the analysis of reconstructed Neogene climate data, we suggest that the intensification of the East Asian winter monsoon during the Quaternary may have dramatically increased seasonality and given rise to a much drier winter in Yunnan. Combined with information on Cedrus fossil records and its seed physiology, we conclude that the intensification of a drier climate after the Late Pliocene may have prevented the survival of Cedrus seedlings, leading to the eventual disappearance of Cedrus in western Yunnan. This study indicates that the topography in southwestern China acted as a vital refuge for many plants during the Quaternary, but that other species gradually disappeared due to the intensification of the monsoonal climate.
Stem and frond remains are described from Middle Pennsylvanian coal balls collected from the Maple Grove Mine, Cayuga, Indiana.
A key for classifying the frass of 24 species of cossid, sesiid, and cerambycid borers describes the presence, texture, form, size, and color of such frass components as excrement pellets, wood chips, finely pulverized wood, granular wood pieces, fibrous shreds, and excelsior-like shreds. Descriptions are based largely upon fresh frass produced by mid- to late-instar larvae. Host tree species, stem diameter, and portion of tree infested are used in conjunction with frass descriptions to identify the borers.
Lower Miocene silicified woods from south‐east Coates Bay, Hukatere Peninsula, Kaipara Harbour, northern New Zealand, were studied using thin sections and scanning electron microscope micrographs. Woods resembling Avicennia, Nothofagus, Leptospermum, Vitex, Quintinia, Eucalyptus, Dacrydium, and other gymnosperms are described. Pyroclastic flows carried woods from a podocarp‐broadleaved forest to mix with Avicennia (mangrove) growing in estuarine conditions close to the site of preservation. The wood was probably alive or recently dead when overwhelmed by a pyroclastic flow. Fossil termite faecal pellets are recorded from borings in Avicennia sp. The hexagonal silicified pellets are similar to those formed by the extant New Zealand dry wood termite Kalotermes brouni. Four other types of borings in the wood and traces of fungal breakdown are described.