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An Early Devonian arthropod fauna from the Windyfield Cherts, Aberdeenshire, Scotland
Abstract and Figures
New terrestrial and freshwater arthropods are described from the Windyfield cherts, a suite of silicified sinters deposited 700m north-east of the Rhynie cherts and part of the same Early Devonian hot-spring complex. The diverse assemblage consists of Heterocrania rhyniensis (Hirst and Maulik, 1926a), here recognized as a euthycarcinoid; scutigeromorph centipede material assigned to Crussolum sp.; the crustacean Lepidocaris; trigonotarbid arachnids; a new arthropod of myriapod affinities named Leverhulmia mariae gen. et sp. nov.; and the distinctively ornamented arthropod cuticle of Rhynimonstrum dunlopi gen. et sp. nov. The Leverhulmia animal preserves gut content identifying it as an early terrestrial detritivore. Abundant coprolites of similar composition and morphology to the gut contents of the euthycarcinoid crowd the matrix. Chert texture, faunal associations, and study of modern analogues strongly suggest that the terrestrial arthropods were ubiquitous Early Devonian forms with no particular special adaptation to localized conditions around the terrestrial hot-spring vents. The aquatic arthropods represent biota from ephemeral cool-water pools in the vicinity of the hot-spring vents.
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... ( Fig. 1P; Jeram et al., 1990). As mentioned below, we also refer to material from the Windyfield Chert (AUGD 12307-12308; Anderson and Trewin, 2003) for morphological details (Fig. 9c-e). ...
... Crussolum was resolved as stem-group Scutigeromorpha in the morphological cladistic analysis of Fernández et al. (2016). Codings were a composite of material described as Crussolum sp. from the Windyfield Chert (Pragian) of the Dryden Flags Formation, Aberdeenshire, Scotland (figured by Anderson and Trewin, 2003), and the one formally named species of the genus, C. crusseratum (Shear et al., 1998), known from isolated and fragmentary legs from the Middle Devonian Gilboa locality, Schoharie County, New York State (Givetian). ...
Fossil age data and molecular sequences are increasingly combined to establish a timescale for the Tree of Life. Arthropods, as the most species-rich and morphologically disparate animal phylum, have received substantial attention, particularly with regard to questions such as the timing of habitat shifts (e.g. terrestrialisation), genome evolution (e.g. gene family duplication and functional evolution), origins of novel characters and behaviours (e.g. wings and flight, venom, silk), biogeography, rate of diversification (e.g. Cambrian explosion, insect coevolution with angiosperms, evolution of crab body plans), and the evolution of arthropod microbiomes. We present herein a series of rigorously vetted calibration fossils for arthropod evolutionary history, taking into account recently published guidelines for best practice in fossil calibration. These are restricted to Palaeozoic and Mesozoic fossils, no deeper than ordinal taxonomic level, nonetheless resulting in 80 fossil calibrations for 102 clades. This work is especially timely owing to the rapid growth of molecular sequence data and the fact that many included fossils have been described within the last five years. This contribution provides a resource for systematists and other biologists interested in deep-time questions in arthropod evolution.
ABBREVIATIONS
AMNH American Museum of Natural History
AMS Australian Museum, Sydney
AUGD University of Aberdeen
BGR Bundesanstalt fur Geowissenschaften und Rohstoffe, Berlin
BMNH The Natural History Museum, London
CNU Key Laboratory of Insect Evolutionary & Environmental Change, Capital Normal University, Beijing
DE Ulster Museum, Belfast
ED Ibaraki University, Mito, Japan
FMNH Field Museum of Natural History
GMCB Geological Museum of China, Beijing
GSC Geological Survey of Canada
IRNSB Institut Royal des Sciences Naturelles de Belgique, Brussels
KSU Kent State University
Ld Musee Fleury, Lodeve, France
LWL Landschaftsverband Westfalen-Lippe-Museum fur Naturkunde, Munster
MACN Museo Argentino de Ciencias Naturales, Buenos Aires
MBA Museum fur Naturkunde, Berlin
MCNA Museo de Ciencias Naturales de Alava, Vitoria-Gasteiz, Alava, Spain
MCZ Museum of Comparative Zoology, Harvard University
MGSB Museo Geologico del Seminario de Barcelona
MN Museu Nacional, Rio de Janeiro
MNHN Museum national d'Histoire naturelle, Paris
NHMUK The Natural History Museum, London
NIGP Nanjing Institute of Geology and Palaeontology
NMS National Museum of Scotland
OUM Oxford University Museum of Natural History
PBM Palaobotanik Munster
PIN Paleontological Institute, Moscow
PRI Paleontological Research Institution, Ithaca
ROM Royal Ontario Museum
SAM South Australian Museum, Adelaide
SM Sedgwick Museum, University of Cambridge
SMNK Staatliches Museum fur Naturkunde, Karlsruhe
SMNS Staatliches Museum fur Naturkunde, Stuttgart
TsGM F.N. Chernyshev Central Geologic Prospecting Research Museum, St. Petersburg
UB University of Bonn
USNM US National Museum of Natural History, Smithsonian Institution
UWGM University of Wisconsin Geology Museum
YKLP Yunnan Key Laboratory for Palaeobiology, Yunnan University
YPM Yale Peabody Museum
ZPAL Institute of Paleobiology, Polish Academy of Sciences, Warsaw.
... Передние лопасти имеют внутренние полукруглые ямки, свидетельствующие о сложном устройстве данного элемента. Он аналогичен мандибулам известных эвтикарциноидов, например, Apankura [ [9]], Heterocrania [ [10]] и Euthycarcinus [11,12]. В медиальной бороздке между этими элементами видно небольшое углубление; это, вероятно, рот. ...
Описывается новое местонахождение Кимильтей, Иркутская область, где в массе захоронились неминерализованные остатки членистоногих. Изотопное U–Pb-датирование детритовых цирконов из слоя с фоссилиями указывает на их кембрийский возраст 491 ± 6 млн лет. Членистоногие в Кимильтей представлены Chasmataspidida, Synziphosurina и Euthycarcinoidea. Две последние группы на Сибирской платформе известны не были, а первая известна лишь в девоне. Иркутские находки попадают в ряд древнейших представителей своих групп, а ареал каждой из них оказывается шире, чем предполагалось. Обращает на себя внимание совместное захоронение этих членистоногих; прежде их не находили вместе. С учетом того, что потомки всех трех групп дали начало наземным линиям хелицеровых и насекомых, то это целая фауна предков наземных членистоногих. Сожительство предков хелицеровых и насекомых указывает, что освоение суши стартовало у них из близких экологических ниш в приливной зоне.
... Haug et al. 2014). The oldest remains of fossil centipedes are fragments of legs of scutigeromorphans from the Silurian (Shear et al. 1998), other fragments are known from the Devonian (Shear et al. 1998;Anderson and Trewin 2003;Haug and Haug 2017). More complete, but still fragmented pieces, are known from the Devonian as cuticle remains of Devonobius delta, which shares some characters with Crateostigmus (Shear and Bonamo 1988). ...
Centipedes are predatory representatives of the group Myriapoda and important components of the soil and leaf-litter fauna. The first pair of trunk appendages is modified into venom-injecting maxillipeds in all centipedes. The number of trunk appendage pairs varies between the different groups of centipedes, from 15 pairs as apparently ancestral (plesiomorphic) condition, up to 191 pairs. The last pair of trunk legs can be used for different tasks in centipedes, e.g. mechano-sensation, defense, or stridulation. Many morphological details are also known from fossil centipedes, but especially the oldest fossils are often fragmentary and the fossil record in general is rather scarce. Especially the late appearance of lithobiomorphans in Cenozoic ambers is notable, though some not formally described lithobiomorph-like specimens from Cretaceous amber from Myanmar have been published. We present here a new specimen from Cretaceous Kachin amber, Myanmar with a lithobiomorph-type of morphology, Lithopendra anjafliessae gen. et sp. nov. The very large ultimate leg appears to have been used for defence and is, in relative proportions, larger than in any known lithobiomorphan, only comparable to that in scolopendromorphans. With this, the specimen presents a mixture of characters, which are in the modern fauna only known from two different centipede groups. We discuss the implications of this new fossil, also concerning events of convergence in this lineage.
... Anterior parts have internal semicircular pits indicating the complex structure of this element. It is similar to the mandibles of known euthycarcinoids, for example, Apankura [9], Heterocrania [10], and Euthycarcinus [11,12]. A small depression is seen in the medial groove between these elements; it is probably a mouth. ...
A new locality Kimiltei, Irkutsk oblast, where unmineralized remains of arthropods were buried en
masse, is described. The U–Pb zircon dating yielded a Cambrian age with the youngest values of 492±4 Ma.
Kimiltei arthropods are represented by Chasmataspidida, Synziphosurina, and Euthycarcinoidea. The last
two groups have not been known in the Siberian Platform, and the first one has been described in this area
only from Devonian rocks. The Irkutsk findings are the most ancient representatives of their groups, and their
distribution areas turn out to be wider than previously known. Their coburial is noteworthy, because the representatives
of these groups have never been found together before. Taking into account the fact that the
descendants of all three groups gave rise to the terrestrial lines of chelicerates and insects, the Kimiltei assemblage
is the entire fauna of the land arthropod ancestors. Their cohabitation is indicative of the fact that they
started their expansion on land from some close ecological niches in the intertidal zone.
... The associated Rhyniognatha hirsti Tillyard, 1928 is now, however, considered a myriapod and not a hexapod (Haug & Haug, 2017). This is in contrast to Leverhulmia mariae Anderson & Trewin, 2003, originally proposed as a myriapod, then a stem hexapod (Fayers & Trewin, 2005). ...
At the third congress of the I.P.S. (International Palaeoentomological Society) in Beijing (2010), Professor Dong Ren, conference organiser, was presented with an author’s copy of the arthropod volume of the Geological Conservation Review of Great Britain, which included fossil insects, and had just been published (Jarzembowski et al., 2010). The purpose of the review, which commenced in the last century before the founding of I.P.S., was essentially to select and document the key sites of British geology, geomorphology and palaeontology with view to geoheritage conservation—including palaeoentomology. The results were collated and published in a series of volumes and, as it subsequently transpired, the arthropod volume was the last one (number 35). The insect part (written by the current author) and other arthropods (by Derek Siveter and Paul Selden) were augmented by general geology and palaeontology contributed by Douglas Palmer. A planned volume with relevant Lower Cretaceous (Wealden) geology was eventually produced instead as a short series of papers by the Geologists’ Association of London (Radley & Allen, 2012b). Geoconservation has featured periodically on the I.P.S. agenda and this paper reflects on the legacy of the GCR study, a decade later, and over a generation after its initiation.
Mixed siliciclastic-carbonate deposits of Lower Devonian (Lochkovian-Emsian) age are exposed in the northeastern part of the Reggane Basin (Touat region). Based on lithological, sedimentological and palaeontological data, these deposits are subdivided into four distinct lithological units: Unit A characterises the Silurian-Devonian transitional zone; Unit B consists of a thick succession of Lochkovian-Pragian age, whereas Units C and D both document Emsian deposits. The studied section exhibits low to moderate ichnodiversity consisting of 14 ichnotaxa: Arenicolites isp., Chondrites isp., Curvolithus multiplex, Lockeia siliquaria, Monomorphichnus isp., Neonereites biserialis, Neonereites uniserialis, Ophiomorpha isp., Palaeophycus isp., Planolites isp., Psilonichnus upsilon, Skolithos linearis, S. verticalis, and Thalassinoides isp. Both Curvolithus multiplex and Psilonichnus upsilon are documented for the first time from Palaeozoic deposits of the Saharan platform. These ichnofaunas are commonly ascribed to the Skolithos and Cruziana ichnofacies. Five distinct ichnoassemblages (Ichnoassemblages A to E) were defined, reflecting palaeoenvironmental changes from the foreshore to upper offshore zones, with frequent storm influence. Ichnofabric analysis of the section studied reveals six levels with different bioturbation intensities showing different palaeoecological conditions in various environmental settings (Monospecific Planolites, Lower combined Skolithos-Arenicolites, Combined SkolithosThalassinoides, Monospecific Skolithos, and Upper combined Skolithos-Arenicolites and Reburrowed Chondrites ichnofabrics). The vertical evolution of these deposits displays the successive deepening and shallowing trends recorded during the Lower Devonian in this area.
Despite the enormous paleobotanical record on different islands of the Antarctic Peninsula, the evidence of insect activity associated with fossilized plants is scarce. Here we report the first evidence of insect-plant interaction from Cretaceous deposits, more precisely from a new locality at the Rip Point area, Nelson Island (Antarctic Peninsula). The macrofossil assemblage includes isolated Nothofagus sp. leaf impressions, a common component of the Antarctic paleoflora. Two hundred leaves were examined, of which 15 showed evidence of insect activity, displaying variations in size, shape, and preservation. Two types of interaction damage, galls and mines, were identified. A single specimen retained a circular scar recognized as galling scar, while meandering tracks were considered mines. These traces of herbivore insect activity, correspond to the oldest known record of this type of interaction of West Antarctica and the oldest record of insect-plant interaction in Nothofagus sp. reported so far.
Animals breaking away from the sea was a revolutionary event in the evolution of life. Arthropods were the earliest metazoans to move onto land, and although a few Silurian freshwater and/or terrestrial arthropods have been found so far, these records are all from Laurussia. Here, we describe a new freshwater arthropod, Maldybulakia saierensis sp. nov., from the western Junggar, northwest China. Evidence from co-occurring spores and body fossils of plants is presented in support of a Silurian (Pridoli) age for this new Maldybulakia species, alongside palaeosalinity data in support of our interpretation of it having lived in a freshwater environment. The discovery of this species brings forward the earliest appearance of the Maldybulakia, previously known from theDevonian of Kazakhstan and eastern Australia, to the late Silurian. It is the oldest body fossil record of a putatively freshwater arthropod outside Laurussia, and greatly expands their
palaeogeographical distribution. In the middle and late Silurian, the discovery of freshwater arthropods on multiple plates/terranes, as well as their morphological diversity during
this period, suggests that arthropods had left the marine environment by the early Silurian or even earlier.
Premise of the Research. The Lower Devonian Rhynie chert preserves abundant microbial fossils that are similar morphologically to cyanobacteria and microscopic algae, but cannot be placed systematically with confidence. Although often morphologically distinct, only a few of these fossils have been described.
Methodology. Thin sections prepared from three different blocks of the Rhynie chert were examined at high magnification in transmitted light. Images of fossils were captured digitally and processed in Adobe Photoshop.
Pivotal Results. More than 150 specimens of Glaphyrobalantium hueberi gen. et sp. nov., a minute vesicle bounded by a prominent envelope and usually containing eight or more spherical bodies, were found. The fossils are similar to certain present-day coccoid cyanobacteria (e.g., Gloeocapsa; Chroococcales) and algae (e.g., Gloeocystis; Sphaeropleales) that occur in microscopic groups of 2–8 (or more) cells surrounded by a colonial sheath, and unicellular algae that reproduce asexually by autospores (e.g., Chlorococcum; Chlamydomonadales), but affinities to protists, fungi, and fungus-like organisms can also not be ruled out.
Conclusions. This discovery expands our knowledge of microbial life in the Rhynie paleoecosystem. Although the affinities remain elusive, it is important to describe fossils like G. hueberi as a first step towards understanding past microbial biodiversity and the roles microorganisms have had as constituents of ancient ecosystems.
Remains of arthropod legs with a pentagonal cross section and serrate margins, found at three Silurian and Devonian sites (Ludford Lane, Wales; Rhynie, Scotland; and Gilboa, New York, USA), are herein attributed to teffestrial scutig-eromorph centipeds. The Silurian and Devonian legs are distinct from each other, from previously described Carboniferous remains, and from mod-em scutigeromorphs, but the general pattern and many of the details of leg construction in these centipeds seems to have been conserved over a 415 million year history. The legs are attributed to a new scutigeromorph genus, Crussolum, placed in a new monobasic family Crussolidae; fairly complete remains from the Devonian of Gil-boa, New York, are assigned to the new species Crussolum crusserratum Shear. The Silurian (Ludford Lane) and Rhynie legs are too poorly known to be given a species epithet, but more than one taxon may be present.
The oldest known spider, from the Devonian (Givetian) of Gilboa, New York, is Attercopus fimbriunguis (Shear, Seldon and Rolfe), parts of which were originally described as a trigonotarbid, possibly of the genus Gelasinotarbus. Previous reports of Devonian spider fossils, from the Lower Emsian of Alken-an-der-Mosel, Germany, and the Pragian of Rhynie, Scotland, are shown to be erroneous identifications. Attercopus is placed as sister-taxon to all living spiders, on the basis of characters of the spinneret and the arrangement of the patella-tibia joint of the walking legs. A cladogram of the relationships of all pulmonate arachnids is presented. A pulmonate arachnid from Gilboa, related to Araneae and Amblypygi, is described as Ecchosis pulchribothrium Selden and Shear, gen. et sp. nov., and additional arachnid material is described. -Authors
The Euthycarcinoidea is a superclass of the arthropod phylum Uniramia and one of the rarest groups of fossil arthropods. Only seven species in five genera have been described, from rocks of Late Carboniferous age in France and the USA, and of Middle Triassic age in France and eastern Australia. Here, a much older euthycarcinoid, from a mixed sequence of fluviatile and aeolian sandstones of probably Late Silurian age in Western Australia, is described as Kalbarria brimmellae gen. et sp. nov. Because of their uniramian affinities, it has previously been suggested that euthycarcinoids may be the closest ancestral relatives of hexapods. Although previous evidence had indicated that the earliest hexapod predated the earliest euthycarcinoid, the discovery of a euthycarcinoid about 120 million years older than the previous oldest record, and predating the oldest known hexapod, provides strong support for the view that the hexapods may have evolved from the Euthycarcinoidea. -from Authors
Well preserved Arthropod-Cuticles have been obtained by HF-maceration from coaly clay- and siltstones of the Rhenish Lower Devonian and the Upper Carboniferous of the Ruhr- and the Saar-basin. They occur is isolated cuticular-sheets, larger isolated body pieces and almost complete animals. Remnants of ostracods and mites are particularly well preserved, the mites being the best up to now described specimens in the fossil record apart from those found in amber. The ostracods show well preserved appendages. Body pieces of small scorpions are also remarkably rich. Chelicerata (presumably Arachnids) yielded cuticular pieces displaying distict sensory organs. The oldest slit sensilla come from the Lower Devonian (Lower Emsian, Waxweiler, Eifel-region), distinct lyriform organs are to be found in the upper Carboniferous. In Sedimentary rocks from the Lower Devonian remnants of eurypterids prevail. In almost all samples different morphologies of hair sensilla were found. The paleophysiological and systematic relationships are discussed. The Arthropods found fall in a size range between 0,5 to 1 mm in complete smaller Arthropods (mites, ostracods), body pieces come from animals with a calculated size of about 5 to 10 mm. This size range falls below that of most hitherto described fossil Arthropods. A method of rebedding larger arthropod fossils from the rock unto an acetate film (here: eurypterids from the Rhenish Lower Devonian) makes the close investigation of cuticular structures possible. Lithological characteristics and preservational/diagenetical effects in view of future prospection are discussed in the cases of all occurrences. The palynodebris investigation of the Lower Devonian Brandschiefer-seams resulted in a more differentiated paleoenvironmental view. Regional and stratigraphical trends in Arthropod contents may be observed in some of the Lower Devonian and Upper Carboniferous occurrences. The palynological methods can be of equal value for Arthropod investigation besides occurrences of phosphatized animals and silicified peats. Due to lithologic characters neccessary it is even more broadly applicable.
Standards have been empirically developed to describe various morphological characters of eurypterids. The standards pertain to the following characters: 1) shape of the prosoma; 2) shape of the metastoma; 3) shape of the eyes; 4) position of the eyes; 5) types of prosomal appendages; 6) types of swimming leg paddles; 7) structure of the doublure; 8) differentiation of the opisthosoma; 9) structure of the genital appendages; 10) shape of the telson; and 11) types of ornamentation.
For the first time, a uniform, standardized taxonomy is proposed for classification and identification of most genera. The taxonomy is based on the observation that most higher taxonomic levels for arthropods are based on the structure and arrangement of the appendages. Details of the taxonomy rely on the morphological standards proposed here.
The order Eurypterida Burmeister, 1843, is here defined by the presence of only six pairs of prosomal appendages, the first pair being the chelicera, the next five pairs being the gnathobasic, uniramus legs. Suborders are characterized by the gross morphology of the chelicera. Superfamilies and families are characterized by the use of a single character complex, specifically the structure and arrangement of the second through sixth pairs of prosomal appendages. Genera are recognized by more specific standards.
A new classification of the order Eurypterida is proposed. Three new superfamilies, Kokomopteroidea, Megalograptoidea, and Brachyopterelloidea, and five new families, Brachyopterellidae, Adelophthalmidae, Lanarkopteridae, Erieopteridae, and Hardieopteridae, are proposed.