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Significant bioinclusions of plants and animals in Southern Gondwana late middle Eocene amber of Anglesea, Victoria. (A to B) Liverworts of the genus Radula (Marchantiophyta: Radulaceae). (C) Two stems with perfectly preserved phyllids or leaf-like structures of mosses of the genus Racopilum (Bryophyta: Racopilaceae). (D) Juvenile individuals of spiders. (E to F) Springtail of the living genus Coecobrya (Entomobryomorpha: Entomobryidae) in two views. (G) A Symphypleona springtail. (H) Light photograph of large piece of yellow amber with two dipterans, Dolichopodidae at left and Ceratopogonidae at right, and at top of image a mite of the living genus Leptus (Arachnida: Acari: Trombidiformes: Erythraeidae). (I) Dipterans of the family Dolichopodidae (long-legged flies) in copula. (J) Worker ant of the living genus Monomorium or a "Monomorium-like" lineage (Hymenoptera: Formicoidea: Formicidae) (see Fig. S2).
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The Northern Hemisphere dominates our knowledge of Mesozoic and Cenozoic fossilized tree resin (amber) with few findings from the high southern paleolatitudes of Southern Pangea and Southern Gondwana. Here we report new Pangean and Gondwana amber occurrences dating from ~230 to 40 Ma from Australia (Late Triassic and Paleogene of Tasmania; Late Cre...
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... ACM amber biota comprises a multitude of animal, plant and fungal groups from a total of 3,291 amber pieces and the most bioinclusion-rich, in situ deposit recorded from Australia (Fig. 4). Fungi include hyphomycetean remains similar to the extant anamorphic genus Monotosporella (Fig. S1). Plant inclusions comprise well preserved non-vascular plants (i.e., bryophytes) of liverworts and mosses (the first in amber in the far south), a trichome, a petal, leaf bract, possible seeds, and a possible megaspore. Significantly, ...
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... well preserved non-vascular plants (i.e., bryophytes) of liverworts and mosses (the first in amber in the far south), a trichome, a petal, leaf bract, possible seeds, and a possible megaspore. Significantly, two species of the liverwort Radula (Radulaceae) have been recognized along with two moss species attributed to Racopilum (Racopilaceae) (Fig. 4A-C). Several arthropods (arachnids and hexapods) have been identified in the ACM amber, which include juvenile spiders in a cluster most likely formed after hatching for safety reasons until the next moult (Fig. 4D) and immature mites belonging to the Trombidiformes (Erythraeidae, Leptus Latreille, 1796). Hexapods are represented by some ...
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... species of the liverwort Radula (Radulaceae) have been recognized along with two moss species attributed to Racopilum (Racopilaceae) (Fig. 4A-C). Several arthropods (arachnids and hexapods) have been identified in the ACM amber, which include juvenile spiders in a cluster most likely formed after hatching for safety reasons until the next moult (Fig. 4D) and immature mites belonging to the Trombidiformes (Erythraeidae, Leptus Latreille, 1796). Hexapods are represented by some collembolans and diverse insect groups, as lepidopterans represented by diverse and abundant, isolated scales (Fig. S1), and flies of the families, Ceratopogonidae (or biting midges; Fig. 4H), Dolichopodidae (or ...
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... reasons until the next moult (Fig. 4D) and immature mites belonging to the Trombidiformes (Erythraeidae, Leptus Latreille, 1796). Hexapods are represented by some collembolans and diverse insect groups, as lepidopterans represented by diverse and abundant, isolated scales (Fig. S1), and flies of the families, Ceratopogonidae (or biting midges; Fig. 4H), Dolichopodidae (or long-legged flies, two of them in copula; Fig. 4I), and Tipulidae (or craneflies). Dolichopodidae have a fossil record extending to the Cretaceous with several records in the Paleogene in amber deposits worldwide. These predatory flies today eat springtails (also recorded in the ACM amber), aphids and a variety of ...
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... to the Trombidiformes (Erythraeidae, Leptus Latreille, 1796). Hexapods are represented by some collembolans and diverse insect groups, as lepidopterans represented by diverse and abundant, isolated scales (Fig. S1), and flies of the families, Ceratopogonidae (or biting midges; Fig. 4H), Dolichopodidae (or long-legged flies, two of them in copula; Fig. 4I), and Tipulidae (or craneflies). Dolichopodidae have a fossil record extending to the Cretaceous with several records in the Paleogene in amber deposits worldwide. These predatory flies today eat springtails (also recorded in the ACM amber), aphids and a variety of larvae in a wide www.nature.com/scientificreports ...
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... Reconstruction with anatomical features of the second fossil ant discovered in Australasia and the oldest from Southern Gondwana. The reconstruction of this worker ant corresponds to a new myrmicine species belonging to the extant genus Monomorium or a "Monomorium-like" lineage discovered in late middle Eocene amber of Anglesea, Victoria (see Fig. 4J). It is based on several specimens (also see Fig. S2), and the body color pattern is conjectural, but aligns with the common pattern found in extant Monomorium ants. Performed using Light-Wave 3D computer graphics program (NewTek; www.newtek.com/lightwave/) (Artist: J.A. Peñas). Blattodea). Two collembolans belong to the so-called ...
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... (Artist: J.A. Peñas). Blattodea). Two collembolans belong to the so-called 'slender springtails' (Hexapoda: Entomobryidae) that, until now, had no fossil record in Australia, but are found elsewhere in the Eocene, including many specimens in Baltic amber. One of these specimens has been classified in the living genus Coecobrya (Fig. 4E,F). The fossil record of springtails is notably poor in the Southern Hemisphere, but Entomobryidae have already been recorded as amber inclusions from the late Oligocene-early Miocene of New Zealand 9 . Exquisitely preserved specimens of collembolans belonging to Symphypleona have been also recovered from the new ACM amber locality (Fig. ...
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... (Fig. 4E,F). The fossil record of springtails is notably poor in the Southern Hemisphere, but Entomobryidae have already been recorded as amber inclusions from the late Oligocene-early Miocene of New Zealand 9 . Exquisitely preserved specimens of collembolans belonging to Symphypleona have been also recovered from the new ACM amber locality (Fig. 4G). Most significant are several winged and worker ants belonging to the myrmicine genus Monomorium, according to the current status of this non-monophyletic genus (they are very similar to the synonymized Chelaner), or a "Monomorium-like" lineage. The specimens exhibit a unique combination of characters indicating that most likely this ...
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... and worker ants belonging to the myrmicine genus Monomorium, according to the current status of this non-monophyletic genus (they are very similar to the synonymized Chelaner), or a "Monomorium-like" lineage. The specimens exhibit a unique combination of characters indicating that most likely this is the fourth known fossil species of Monomorium (Figs. 4J, 5, 6 and S2) and the first recorded in the Southern Hemisphere 45 . In the Recent biota, Monomorium mostly inhabits the Old World, particularly the tropics, and Australia is considered one of the two main centers of speciation. These new fossil ants are the oldest from Southern Gondwana and unveil the antiquity of many elements in the ...
Citations
... Moreover, insects trapped in fossil resins attracted frequently predators and scavengers. Often, ambers also contain not only trapped pollen, or spores but also small fragments of plant tissues (e.g., Solórzano-Kraemer, 2010;Wang et al., 2014;Sadowski et al., 2017;Stilwell et al., 2020). Therefore, ambers are "time capsules" that may contain remains of organisms from past ecosystems in intact, morphologically well-preserved forms. ...
... A modern species -Agathis australis (kauri tree) was often investigated and compared to extant plants of the same family . The major amber occurrences in Australia and Oceania are listed in Table 6 Stilwell et al., 2020). The chemical composition of the ambers, however, has not been investigated so far. ...
... Ambers were also found in the Paleogene sediments in Australia, Tasmania, and New Zealand (Victorian Paleocene amber, Pomahaka amber; nos. 71, 75; Lyons et al., 2009;Stilwell et al., 2020). The botanical source of the ambers has not been fully elucidated yet. ...
Fossil resins, also known under the name “ambers”, are formed from exudates of gymnosperm and angiosperm trees in Earth's sedimentary rocks. Up to day, more than 167 fossil resin occurrences and major deposits have been reported in literature. The scientists are mainly focused on fossiliferous ambers, which contain numerous fossil inclusions. However, the geochemical studies, especially those that are aimed at elucidation of the chemical composition of the fossil resins are scantily reported in literature. The chemical composition of ambers is very important for establishing an evolutionary link between ancient plants and their present form. Here, we present seventy-eight major occurrences of fossil resins together with their geochemical, botanical, and geographical data that have been published up to now. Most of them are from North America, Europe, and Asia; however, the occurrences from South America, Africa, Australia, and Oceania are also discussed. The examined ambers have been found in the late Carboniferous or younger sediments. In most cases, the sediments were formed in a close-to-the-shore or swamp-based environment. In many cases, the fossil resins were likely transferred and redeposited in younger sediments during various environmental and geological processes.
... 12 From the Cretaceous onward, ambers start to provide a window to the larval forms of Holometabola in the past in an almost life-like manner. While there are older ambers, [13][14][15][16] these have so far not included holometabolan larvae. Among the different Cretaceous ambers, Kachin amber, found in Myanmar and being around 100 million years old, has been especially productive in providing examples of holometabolan larvae 17,18 that possess different types of anti-predator structures or allow to infer anti-predator types of behavior. ...
Holometabolan larvae are a major part of the animal biomass and an important food source for many animals. Many larvae evolved anti-predator strategies and some of these can even be recognized in fossils. A Lagerstätte known for well-preserved holometabolan larvae is the approximately 100-million-year-old Kachin amber from Myanmar. Fossils can not only allow to identify structural defensive specializations, but also lifestyle and even behavioral aspects. We review here the different defensive strategies employed by various holometabolan larvae found in Kachin amber, also reporting new cases of a leaf-mining hymenopteran caterpillar and a hangingfly caterpillar with extensive spines. This overview demonstrates that already 100 million years ago many modern strategies had already evolved in multiple lineages, but also reveals some cases of now extinct strategies. The repetitive independent evolution of similar strategies in distantly related lineages indicates that several strategies evolved convergently as a result of similar selective pressures.
... Cretaceous amber records are sparse from the Southern Hemisphere, being reported from South America, Australasia and Africa (Petrulevicǐus and Iglesias 2018;Bouju and Perrichot 2020;Seyfullah et al. 2020;Stilwell et al. 2020). The last is the only to yield arthropod inclusions to date (Table 1). ...
Amber first became relatively abundant and widespread in the geological record during the Cretaceous period. It originated often as copious resin production by a variety of incompletely understood coniferous trees, generally under humid climates, but not excluding seasonal aridity. Study of insect and other organic inclusions only commenced in the twentieth century, but has expanded considerably since then, with several thousand taxa now described. Cretaceous amber insects can be exquisitely preserved in three dimensions, although tend to be biased towards smaller individuals that lived in the local forest environment. They are therefore complemented by the Cretaceous rock record, which sampled larger insects and other habitats. As well as the fine morphological detail exhibited by the amber insect inclusions, various behaviours and interactions unlikely to be otherwise preserved can be found frozen in time, such as brooding behaviour and the entrapment of insects in spiders' webs. Insects in amber also provide important information about evolutionary changes over the course of the Cretaceous, including the rise of eusociality and angiosperm pollination.
... Consequently, the migration of these psocid families from south-eastern Asia to the Burma Terrane cannot be ruled out. Therefore, the discovery of new troctomorphans in African (Bouju and Perrichot, 2020) or Australian (Stilwell et al., 2020) Cretaceous ambers would help to determine the evolutionary origin of these groups. Interestingly, although the Iberia Island was geographically close to Laurasia, the psocid fauna from Spanish amber seems to be more similar to that from Lebanese and Burmese ambers (Table 1). ...
Psocids, commonly known as barklice, are insects belonging to the order Psocodea, together with the parasitic lice. They usually inhabit forest litter or the bark of tree trunks and branches, showing grazing herbivorous or detritivorous feeding habits. The Cretaceous psocid record is diverse, containing more than 70 described species. Here, we present new psocids (Troctomorpha and possible Psocomorpha) from two Spanish amber outcrops, both Albian (Lower Cretaceous): El Soplao and Ariño. We describe the two new species Azarpsocus anjana Álvarez-Parra and Nel sp. nov. (Manicapsocidae) and Burmacompsocus ojancano Álvarez-Parra and Nel sp. nov. (Compsocidae), and the morphotype Ariño that might belong to Psocomorpha. We discuss the taxonomic placement of the studied specimens and comment on the palaeobiogeography of the Cretaceous psocids. The Cretaceous barklice fauna from Iberia shows more similarities with that from Lebanese (Barremian) and Burmese (Cenomanian) ambers than with that from the palaeogeographically closer French amber (Cenomanian). This finding has been also reported in other insect groups from Spanish amber, and might be the consequence of several factors, such as the palaeoenvironmental conditions of the ecosystems, the palaeogeographical position of the Iberia Island during the latest Jurassic, closer to Gondwana than to Laurasia, and oceanic currents that facilitated or hindered the displacement of insect fauna between continental masses.
... In the past, researchers have mainly studied amber fossils with the aim of discovering new insect species, leaving nematodes -including mermithids -and other potential hosts undersampled or understudied (Schmidt et al., 2010;Stilwell et al., 2020;Košulič and Mašová, Parasitic Relationships | Trapped in time 2019). Further investigations will require systematic sampling and global collaboration among all relevant communities and research institutions, along with sufficient funding. ...
Analysis of specimens preserved in amber from the Cretaceous period suggests that nematodes changed their host preference towards insects with a complete metamorphosis more recently.
... An extensive study of dispersed cuticles suggests a much higher overall species diversity (Pole 2007). Last, amber, probably originating from Araucariaceae (Coward et al. 2018), with arthropod, fungal, and plant inclusions (Stilwell et al. 2020) adds to the small number of described fossils. ...
... However, findings of fossil resins in the Paleozoic sediments are regarded as extremely scarce. Similarly, the Late Triassic resin specimens were described only in a few locations, for example, Italy (Gianolla et al., 1998;Roghi et al., 2006Roghi et al., , 2022, Spain (Peñalver and Delclòs, 2010), Tasmania (Stilwell et al., 2020), and Lesotho (Ansorge, 2007). Most likely, these findings reflect the Late Triassic (Carnian) pluvial event, responsible for the global warming which prompted trees to increased resin exudation (Seyfullah et al., 2018;Dal Corso et al., 2020). ...
Fossil resins are the one of several fossil tree products, together with leaves, wood, seeds, or pollen. As an emanation of organic matter, resins are made of organic compounds, predominantly terpenes. Terpenes are broadly identified from various living organisms such as higher plants, fungi, or bacteria. In fossil resins, these compounds occur in several forms-usually mono-, sesqui-, di-, and tri-terpenes. Some terpenes present in fossil resins provide information about their ancestors and the paleoenvironmental conditions during the resin expulsion. These compounds, known as biomarkers, preserve the original shapes of molecules or occur as altered structures. Due to the chromatographic methods, insight into the chemical structure of fossil resins is enabled and biomarkers might be detected. In this study, information about the state-of-the-art chromatographic research concerning fossil resins was gathered. The biomarkers found in fossil resins were divided into compounds providing chemotaxonomic (botanical biomarkers) and environmental information (other biomarkers). Botanical biomarkers provide high utility in paleobotanical studies. The initial identification of a source tree, gymnosperm, or angiosperm is possible, due to certain chemical patterns of these types. Moreover, in chemotaxonomic studies, the restriction to more specific taxonomic levels, especially to the level of family, is usually possible, thanks to the occurrence of specific biomarkers. Many terpenes that are known to have an ecological function in extant plants are also found in fossil resins; however, their presence does not clearly indicate the ecological properties of trees due to the advanced alterations of the chemical structure. However, some specific environmental bio-markers found in fossil resins (e.g. succinic acid or rosane) might provide information about the paleoenvir-onment during the resin deposition.
... Grimaldi (1990) refers to a macrocerine keroplatid and a couple of specimens that may correspond to mycetophilids from the Crato Formation, but they are not well preserved and could not be described. There are big hopes for what is going to be found in the Mesozoic deposits from Australia (Stilwell et al., 2020) as a means to understand the evolution of the Gondwanan insect fauna, including the fungus gnats. At this stage, the leiine species described in this paper is the only fossil of the subfamily to be formally described from Cretaceous Gondwanan deposits. ...
A fossil Mycetophilidae from the Aptian Crato Formation—Cretomanota gondwanica gen. nov., sp. nov.—is described, which is the first mycetophilid from the Crato Formation and corresponds to the oldest known fossil leiine and only the second Gondwanan fossil mycetophilid described so far. Cretomanota gondwanica and both species of Alavamanota Blagoderov and Arillo were added as terminals to the data matrix of a general phylogenetic analysis of the Mycetophilidae, and both fit into the Leiinae. Alavamanota is monophyletic, sister to the clade composed by Cretomanota and the extant genus Manota Williston. The biology of the extant members of this fungivorous family corroborates the reconstruction of the Crato palaeoenvironment as including woodlands with humid habitats and microhabitats. The presence of a Cretaceous member of the tribe Manotini at low latitudes in South America reinforces the hypothesis that the clade with all manotines except Leiella Edwards corresponds to a Lower Cretaceous offshoot from a group in southern Gondwana expanding its distribution to more northern areas into the Gondwana and into Laurasia.
... Ancient tree resin provides fossils in an exceptional preservation state (Grimaldi, 1996;Taylor et al., 2009;Penney, 2010;Ragazzi & Schmidt, 2011). These amber inclusions represent an important source of taxonomic, ecological, or even environmental information, which lead to a better understanding of the evolutionary and geographical history of certain taxa and a better knowledge of the palaeoenvironmental and geological context of a region (Penney, 2010;Wang et al., 2014;Schmidt et al., 2018;Zheng et al., 2018;Stilwell et al., 2020). ...
Amber is renowned for the exceptional preservation state of its inclusions, allowing detailed morphological analysis and providing relevant environmental, palaeoecological, geographical, and geological information. Amber deposits predominantly known from North America, Europe, and Asia, are considered to be rare on the continents that formed Gondwana. The recent discovery of fossiliferous amber deposits in Ethiopia therefore provides an inimitable opportunity to close gaps in the fossil record of African terrestrial biota, and to study organisms otherwise rare in the fossil record. Here we show that diverse cryptogams are preserved in highest fidelity in Miocene Ethiopian amber. We describe gametophyte fragments of four liverworts: Thysananthus aethiopicus sp. nov. (Porellales, Lejeuneaceae), Lejeunea abyssinicoides sp. nov. (Porellales, Lejeuneaceae), Frullania shewanensis sp. nov. (Porellales, Frullaniaceae), and Frullania palaeoafricana sp. nov. (Porellales, Frullaniaceae). Furthermore, we describe a pleurocarpous moss of the extant genus Isopterygium (Hypnales, Pylaisiadelphaceae) and a lichen representing the order Lecanorales. These new specimens represent the first amber fossils of liverworts, mosses, and lichens from the African continent and render Ethiopian amber one of the few worldwide amber deposits preserving bryophytes (mosses and liverworts) or lichens. Fossil species of Thysananthus were recorded in Eocene Baltic and Oligocene Bitterfeld as well as Miocene Dominican and probably also Miocene Mexican ambers. Fossils which can unequivocally be assigned to Lejeunea have only been found in Dominican amber, so far. Neotropical ambers contain only one taxon of Frullania to date, while the genus is most diverse in Baltic, Bitterfeld, and Rovno ambers, formed in temperate regions. The new fossils support a tropical to subtropical origin of Ethiopian amber. The new African liverwort fossils are included in an updated list of leafy liverworts described from worldwide Cenozoic ambers to date. This article is protected by copyright. All rights reserved.
... Carnian amber has yielded the oldest inclusions in amber in the fossil record. Although the search for inclusions also in other Triassic ambers is promising (Stilwell et al., 2020), a substantial amount is only found from the Heiligkreuz Fm., in the Italian Dolomites, so far. Also, here a large volume of small amber droplets is preserved in paleosols (Preto and Hinnov, 2003;Schmidt et al., 2006;Breda et al., 2009;Schmidt et al., 2012). ...
... For example, the amber found inside seeds in South Africa (Ansorge, 2007), in the Molteno Formation (Karoo Basin) is slightly younger with respect to the CPE; it is probably Tuvalian in age (Langer, 2005), whereas the amber from the Triassic Blue Mesa Member of the Chinle Formation of Arizona (United States) (Litwin and Ash, 1991) is most likely younger, potentially even Norian in age (Seyfullah et al., 2018a). In the Fingal Valley Upper Triassic Coal Measures of the Upper Parmeener Supergroup, Tasmania, small amounts of ambers with very few miniscule inclusions are reported (Stilwell et al., 2020). ...
The Carnian Pluvial Episode (CPE) has been recognized as a time of plant radiations and originations, likely related to observed swift changes from xerophytic to more hygrophytic floras. This suggests that the increasing humidity causally resulting from LIP volcanism was the trigger for these changes in the terrestrial realm. Understanding the cause and effects of the CPE on the plant realm, requires study of well-preserved floras that are precisely aligned with the CPE. We therefore focus on the best age-constrained section within the CPE for the terrestrial to marginal marine environment to understand the floristic composition at the early CPE. This is found in the Dolomites, Italy, and is remarkable for the preservation of the oldest fossiliferous amber found in the rock record. An integrated study of palynomorphs and macro-remains related to the conifer families of the fossil resin bearing level brings together the floral components from this section. This observed mixture of different taxa of extinct and modern conifer families underlines firmly the effects of the LIP-induced CPE on the evolution and radiation of conifers.
