Joseph BottingGuest Scientist at Nanjing Institute of Geology and Palaeontology · Living in Llandrindod, Wales
Joseph Botting
PhD
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Introduction
Living and working in the Ordovician Builth Inlier of Wales, and trying to reconstruct in as much detail as possible the ecosystem development in the middle of the GOBE (now also expanding into the surrounding Silurian rocks and their rich echinoderm faunas). I'm also working on new exceptionally-preserved biotas, mostly in the Ordovician of Wales (watch this space...); and of course on the early evolution of sponges, worldwide.
Publications
Publications (113)
Modern poriferans are classified into four classes—Calcarea, Demospongiae, Hexactinellida and Homoscleromorpha—the recognition of which in fossil specimens almost exclusively relies on spicule morphology and arrangement. Early fossil representatives of the phylum Porifera are morphologically diverse, and many of them problematically display charact...
Sponges are among the most common and diverse Cambrian fossils. However, the origin of the crown groups of the extant sponge classes constitutes the principal problem in the understanding of the evolution of these lower metazoans. New intact spiculate sponges from the lower Cambrian Stage 3–Stage 4 Sinsk Lagerstätte of the Siberian Platform enable...
Echiurans (spoonworms) are a very distinctive group of polychaete annelids that had long been considered to constitute a separate phylum. Their fossil record is extremely limited, although trace fossils that have been suggested to be attributable to them date back as far as the Cambrian Period. The oldest body fossils are from the Carboniferous Maz...
Siliceous marine ecosystems play a critical role in shaping the Earth's climate system by influencing rates of organic carbon burial and marine authigenic clay formation (i.e., reverse weathering). The ecological demise of silicifying organisms associated with the Permian‐Triassic mass extinction is postulated to have elevated marine authigenic cla...
Siliceous marine ecosystems play a critical role in climate regulation and the severe impact of the Permian-Triassic mass extinction on silica-secreting animals may have contributed to a sustained greenhouse climate during the Early Triassic, via decreased export of organic carbon burial and increased reverse weathering rates. Yet, our understandin...
Siliceous marine ecosystems play a critical role on the Earth’s climate system through its influence on organic carbon burial and rates of marine authigenic clay formation (i.e. reverse weathering). The ecological demise of silicifying organisms associated with the Permian-Triassic mass extinction is postulated to have elevated rates of marine auth...
Cyathophycus is one of the most widespread reticulosan sponges in Ordovician to Devonian rocks, but was mostly limited to the Iapetus region during the Ordovician and Silurian periods. Having an unfused skeleton, these sponges require an element of exceptional preservation (abrupt burial as a minimum) to enter the fossil record. Most species are a...
An early form of eurypterids (Chelicerata), Archopterus anjiensis n. gen. n. sp., is described from the uppermost Ordovician Wenchang Formation of Anji County, Zhejiang Province, South China. It is the earliest record of eurypterids in China and likely the oldest representative of the Adelophthalmidae. The species, represented by a single specimen,...
Burgess Shale-type faunas are critical to our understanding of animal evolution during the Cambrian, giving an unrivalled view of the morphology of ancient organisms and the ecology of the earliest animal-dominated communities. Rare examples in Lower Ordovician strata such as the Fezouata Biota illustrate the subsequent evolution of ecosystems but...
A crucial step in the evolution of Euarthropoda (chelicerates, myriapods, pancrustaceans) was the transition between fossil groups that possessed frontal appendages innervated by the first segment of the brain (protocerebrum), and living groups with a protocerebral labrum and paired appendages innervated by the second brain segment (deutocerebrum)....
The Cambrian strata of western central Utah have yielded four Miaolingian exceptionally preserved biotas, which are dominated by arthropods and sponges. Arthropods have been continuously studied by various specialists since the first discoveries of soft-bodied fossils in these deposits, but sponges were single-handedly described by J. K. Rigby in s...
Recent studies have described an oceanic anoxic event during the Hirnantian (HOAE) and linked this event to the Late Ordovician mass extinction (LOME). However, the extent and duration of the HOAE remain under debate, as do questions about how oceanic anoxia impacted marine ecosystems. For this study, we investigated two previously unstudied sectio...
he deep-sea hexactinellid sponge Euplectella is an iconic living genus with no known fossil record. The family Euplectellidae (which includes a subset of genera that share the common name ‘Venus’ flower basket’) has a very sparse record from the Middle Cretaceous onwards, and an estimated crown-group origin at around 300 Ma based on molecular clock...
The Anji Biota is a unique deposit with exceptionally preserved sponges and other organisms in deep-water, uppermost Ordovician mudstone of Zhejiang Province, China. The sponge fossils preserve remarkable details including carbonaceous preservation of soft tissues and axial filaments of spicules, but the taphonomy of the deposit has not previously...
Chitinozoans are most commonly known to occur as isolated vesicles, and less commonly (but still regularly) in chains i.e. linear catenary structures. Chitinozoan clusters have been little studied, but are critical to the question of the biological affinity of chitinozoans. Bedding-plane assemblages and acid-digestion residues from Ordovician rocks...
Modern hexactinellid sponges are diverse, but almost exclusively deep-marine organisms with a very intermittent fossil record. Aside from the fused skeletons of hexactinosan lineages (which are also exceptionally rare in Palaeozoic sediments), identifying other families is challenging due to the microscopic nature of many diagnostic characters, and...
A new genus and species of planktic graptolite, Anjigraptus wangi gen. et sp. nov., is described from the Upper Ordovician (Hirnantian) sponge-dominated Anji Biota of Zhejiang Province, China. The new genus has a biserial proximal portion and two uniserial stipes distally, the same tubarium form as Dicranograptus. The new genus has probable pattern...
The Hirnantian glaciation, which was the first in the Phanerozoic to occur with elevated CO2 levels, caused severe mass extinctions and disrupted biogeochemical cycling in the latest Ordovician. Previous research that aimed at exploring the triggers and consequences (both abiotic and biotic) of this glacial event have largely been based on condense...
The protomonaxonid sponges are acontroversial early group with supposed ties to the demosponges, but the group also possesses features that imply amuch earlier-branching position in sponge evolution. Anew species, Choiaella hexactinophora sp. nov., shows atypical protomonaxonid body plan but also contains small hexactin-based spicules, askeletal el...
Originally considered as large, solely Cambrian apex predators, Radiodonta—a clade of stem-group euarthropods including Anomalocaris—now comprises a diverse group of predators, sediment sifters and filter feeders. These animals are only known from deposits preserving non-biomineralized material, with radiodonts often the first and/or only taxa know...
The latest Ordovician to earliest Silurian graptolite Avitograptus avitus is important in the biostratigraphy of the Ordovician–Silurian boundary interval. Two additional species of Avitograptus are described from the sponge-dominated Anji Biota of the Upper Ordovician Wenchang Formation ( Metabolograptus persculptus Biozone) of Zhejiang Province,...
The Anji Biota of Zhejiang Province, South China, is an exceptionally preserved, sponge-dominated fauna from the latest Ordovician interval, representing a deep-water environment and containing more than 100 sponge species. Herein a complex of two common species that together dominate the deepest-water sponge assemblages within the sequence are des...
Early Palaeozoic crinoids are known only patchily from the British Isles except for accumulations at starfish beds. A single, exquisitely preserved crinoid is reported from the Telychian (Llandovery, Silurian) of the Pysgotwr Grits Formation of the Llangurig area, Powys, mid-Wales. This sedimentary succession is turbiditic in origin and poorly foss...
A number of putative annelid worms have been described from Ordovician strata, and these records are included in large-scale compilations of paleontological data. If these fossils are worms, they may yield important phylogenetic information; conversely, if they are not worms, they should not be included in large-scale databases. In either case, res...
Despite the recent discovery of abundant and diverse latest Ordovician sponges in China, early Silurian sponges are generally scarce, and non-lithistid, spiculate sponges remain poorly known. This is particularly true outside of a few well-studied regions, and monographic biases strongly influence the published record of the group. A new, well-pres...
Sponges (Porifera), as one of the earliest-branching animal phyla, are crucial for understanding early metazoan phylogeny. Recent studies of Lower Palaeozoic sponges have revealed a variety of character states and combinations unknown in extant taxa, challenging our views of early sponge morphology. The Herefordshire Konservat–Lagerstätte yields an...
Protomonaxonid sponges are a major group of Cambrian and Ordovician fossils in exceptionally preserved (especially Burgess Shale-type)faunas, but are rare thereafter. Rare examples of apparent surviving lineages are known from the late Palaeozoic and Mesozoic, but by this time more derived groups of sponges have generally displaced them in at least...
The Pentland Hills sponge fauna (Llandovery, Telychian) consists of an unusual, aberrant assemblage, but of low diversity. A new specimen of a unique sponge, Eoghanospongia carlinslowpensis gen. et sp. nov., is described from the classic locality of R82. The mushroom-shaped, probably stalked body (peduncle attachment to body not exposed) resembles...
Agglutinated tubes are produced by a variety of marine organisms. Such tubes will readily break down after the death of the producer, and hence are likely to be found only in deposits with rapid burial and/or exceptional preservation. Here we document agglutinated tubes from four localities of Early Ordovician age. The Lower Ordovician (upper Trema...
The protomonaxonid sponge Pirania is an iconic member of the Burgess Shale community, but recent discoveries show that piraniids are morphologically diverse and also occur in Ordovician strata. Here we establish three new piraniid genera on the basis of Ordovician material from China, Wales, Scotland and Morocco: Auraeopirania gen. nov., Cannapiran...
A variety of unrelated organisms produce agglutinated tubes as dwelling structures, and the habit has persisted since the Neoproterozoic. Onuphionella , an agglutinated tube composed of mica flakes, has been found in lower Cambrian strata on several palaeocontinents. In this paper, a new species of the genus, Onuphionella corusca sp. nov., is descr...
Sponges are believed by many researchers to be the earliest living animal group, but there is conflicting evidence for the timing of their origin. A molecular fossil discovery supports the contention that sponges appeared very early, but starkly contradicts the body fossil record.
The crinoid fauna of the Late Ordovician (Sandbian and Katian) of Morocco is sparse, but diverse. The one previously described species, Rosfacrinus robustus Le Menn & Spjeldnaes, 1996 , is now known to be part of an assemblage of at least 10 species, including the new taxa Iocrinus ouzammoui sp. nov. (Disparida), Superlininicrinus advorsa gen. et s...
The living hexactinellid sponge family Rossellidae is characterised by hypodermal pentactine prostalia and reduction of hexactins to diactins in irregular choanosomal spicule strands. The fossil record of this family extends back to the Upper Cretaceous (˜88 Ma), consistent with molecular-clock analyses that predict an origin at approximately 150 M...
Offshore benthic communities were sparse after the end-Ordovician mass extinction. The recent discovery of diverse, abundant sponges in south-eastern China (Anhui and Zhejiang provinces) that flourished during the Hirnantian (Late Ordovician) post-extinction interval raises questions over the extent and ecological significance of the sponge communi...
Non-lithistid spiculate sponges were major components of Cambrian and Ordovician communities, but currently there is limited understanding of how sponge distribution patterns developed during the Cambrian Explosion and the Great Ordovician Biodiversification Event (GOBE). We compiled data on Cambrian and Ordovician sponge occurrences from the liter...
A small number of organisms create exoskeletons not through biomineralisation, but rather through agglutination: binding sedimentary particles from their environment in a cement or organic matrix. Most agglutinating organisms use any and all materials available, but a small proportion show greater selectivity, for particle size or particle kind.
He...
The two major extant groups of siliceous sponges, Demospongiae and Hexactinellida, are generally regarded as sister groups forming the clade Silicea, although the nature of their last common ancestor is uncertain. The fossil record contains a diverse range of basal demosponges that appear to have evolved from hexactine-bearing reticulosan ancestors...
Sponges have been a major part of marine ecosystems, in both shallow and deep water, from the time of the earliest animal communities. The great shifts in climate that have occurred over the past 541 million years have affected all organisms, including sponges. Although patchy knowledge of the sponge fossil record hinders recognition of trends, som...
A small assemblage of extremely well-preserved fragments of new sponges has been discovered in calcipelites of the middle Cambrian El Mogallón Formation in the Cerro El Mogallón section, near Arivechi in eastern Sonora, Mexico. The assemblage includes two new reticulosan species referred to Ratcliffespongia arivechensis sp. nov. and Valospongia son...
A small assemblage of extremely well-preserved fragments of new sponges has been discovered in calcipelites of the middle Cambrian El Mogallón Formation in the Cerro El Mogallón section, near Arivechi in eastern Sonora, Mexico. The assemblage includes two new reticulosan species referred to Ratcliffespongia arivechensis sp. nov. and Valospongia son...
Sponges have been a major part of marine ecosystems, in both shallow and deep water, from the time of the earliest animal communities. The great shifts in climate that have occurred over the past 541 million years have affected all organisms, including sponges. Although patchy knowledge of the sponge fossil record hinders recognition of trends, som...
Sponges are one of the critical groups in understanding the early evolution of animals. Traditional views of these relationships are currently being challenged by molecular data, but the debate has so far made little use of recent palaeontological advances that provide an independent perspective on deep sponge evolution. This review summarises the...
The Late Ordovician (Hirnantian, approximately 445 million years ago) extinction event was among the largest known, with 85% species loss [1]. Post-extinction survival faunas are invariably low diversity, especially benthic communities [2], but ecological structure was restored relatively rapidly [1]. This pattern, however, reflects organisms with...
In the wake of the end-Permian mass extinction, the Early Triassic (~251.9 to 247 million years ago) is portrayed as an environmentally unstable interval characterized by several biotic crises and heavily depauperate marine benthic ecosystems. We describe a new fossil assemblage —the Paris Biota— from the earliest Spathian (middle Olenekian, ~250.6...
Supplementary material (text, figures and tables) to: Brayard et al. 2017 - Science Advances 3, e1602159.
The Late Ordovician (Hirnantian, approximately 445 million years ago) extinction event was among the largest known, with 85% species loss [ 1 ]. Post-extinction survival faunas are invariably low diversity, especially benthic communities [ 2 ], but ecological structure was restored relatively rapidly [ 1 ]. This pattern, however, reflects organisms...
The Sirius Passet Biota of the Buen Formation in North Greenland is one of the key Burgess Shale-type faunas, as it represents the only diverse early Cambrian (Stage 2, Series 3) biota from Laurentia. The sponges are one of the major components of the fauna, although they appear to be much less abundant than arthropods, based on the available colle...
Sponges form a significant component of the diversity of the Tremadocian (Early Ordovician) Fezouata Biota in the Anti-Atlas of Morocco, but are distributed intermittently and have so far received only limited attention. New material reveals a high diversity of undescribed taxa (a total of at least 27 species), including numerous representatives of...
Exceptional preservation of non-biomineralized arthropods, sponges and vermiform taxa occurs in the Darriwilian (Middle Ordovician) Llanfallteg Formation of South Wales, UK. The assemblage contains elements typical of Ordovician communities juxtaposed with those more commonly associated with Cambrian Konservat-Lagerstätten. This assemblage is prese...
A variety of echinoderm ossicles have been discovered in the middle to upper Cambrian rocks of the North China Craton. Material was collected from two localities: Shuangqiao (SQ) section (Paibian) in Hebei Province and Linyi (LY) (Guzhangian?–Paibian?) in Shandong Province. Six types of ossicles can be recognized, including rounded thecal plates, a...
The Cambrian Burgess Shale-type biotas form a globally consistent ecosystem, usually dominated by arthropods. Elements of these communities continued into the Early Ordovician at high latitude, but our understanding of ecological changes during the Great Ordovician Biodiversification Event (GOBE) is currently limited by the paucity of Ordovician ex...
The Builth Inlier contains a sequence of siliciclastic and volcanic rocks of Middle to Late Ordovician age, representing a range of water depths in a volcanic island setting. Sponges are preserved at sites representing all preserved palaeoenvironments; they are a major part of the fauna at many localities and are dominant in abundance and in divers...
Pelmatozoans diversified primarily during the Middle and Late Ordovician Period, with Early Ordovician records being much more limited, resulting in many gaps in our knowledge of the early stages of their diversification. Dissociated pelmatozoan ossicles have been found abundantly in one section in the Tonggao Formation (Tetragraptus approximatus B...
Calibration of the divergence times of sponge lineages and understanding of their phylogenetic history are hampered by the difficulty in recognizing crown versus stem groups in the fossil record. A new specimen from the lower Cambrian (Series 2, Stage 3; approximately 515 Ma) Sirius Passet Biota of North Greenland has yielded a diagnostic spicule a...
A diverse assemblage of silicified sponges, including orchocladines, rhizomorines, stromatoporoids, hexactinellids and non-lithistid demosponges, has been collected from scree accumulations in a restricted area along the western coast of the Island of Gotland, Sweden. The assemblage comprises 29 species in 20 genera, 18 species of which have not pr...
There is currently little consensus on the branching order and phyletic status of the oldest metazoan
groups, but sponges are widely believed to be the earliest-branching living metazoans. Porifera are thought to have diverged before the emergence of developmental characters typical of Eumetazoa, such as well-defined symmetry; extant sponges show r...
Sponges are probably the earliest branching animals, and their fossil record dates back to the Precambrian. Identifying their skeletal structure and composition is thus a crucial step in improving our understanding of the early evolution of metazoans. Here, we present the discovery of 505-million-year-old chitin, found in exceptionally well preserv...
Abstract: The Cambrian, Ordovician and Silurian distributions of non-stromatoporoid sponges are reviewed. The earliest Cambrian
faunas contain mostly hexactinellids, with protomonaxonids dominating middle Cambrian assemblages. There are no obvious palaeobiogeographical
patterns, with many genera being found widely. Vauxiids, lithistids and heteract...
A new locality exposing Tremadocian (Early Ordovician) graptolitic shales in Hunan Province, South China, has yielded an exceptionally well-preserved annulated worm. This palaeoscolecidan is described as Waflascolex changdensis gen. et sp. nov. and reveals extremely fine detail of the cuticle organization and plating array. The new taxon is charact...
Conulariella sp. and possibly one other conulariid species occur in close association with Sphenothallus sp. in the lower part of the early Floian Tonggao Formation near the town of Sandu, Guizhou Province, China. This is the only known occurrence of Conulariella in Early Ordovician rocks outside of Bohemia (Perunican terrane), and also the first r...
Exceptionally well‐preserved specimens of the reticulosan sponge Cyathophycus loydelli from the Sandbian (Late Ordovician) Llanfawr Mudstones Formation of Llandrindod, Waes, UK, have been examined using scanning electron microscopy (SEM). The specimens include exquisitely detailed pyritized spicules, and granular pyritization of surrounding soft ti...
The Builth Inlier of central Wales exposes a highly fossiliferous Middle to Late Ordovician (Darriwilian to basal Sandbian) siliciclastic succession in a volcanic, back-arc basin setting. Articulated echinoderm faunas are preserved in a range of paleoenvironments, together with widespread, dissociated ossicles. These have enabled a reconstruction o...
My Fezouata data observation have been cited in this publication via communications to the authors.
The Protomonaxonida consist of a heterogeneous group of early fossil sponges traditionally assigned to the demosponges. However, an affinity to the hexactinellid-like Reticulosa has also been suggested, and their relationships are potentially criti...
Abstract: The Cambrian, Ordovician and Silurian distributions of non-stromatoporoid sponges are reviewed. The earliest Cambrian
faunas contain mostly hexactinellids, with protomonaxonids dominating middle Cambrian assemblages. There are no obvious palaeobiogeographical
patterns, with many genera being found widely. Vauxiids, lithistids and heteract...
The comment by Smith et al. rejects a previous interpretation of Middle Ordovician fossils as holothurians. This rejection is based on the use of inappropriate techniques to study the fossils, and insufficient consideration of their taphonomy. The fossils can be shown not to be sponges (the suggested alternative interpretation), and despite the tap...
A new sponge, Heteractenigma yui gen. et sp. nov., is described from the Corymbograptus deflexus Biozone (Floian) of the Ningkuo Formation in western Zhejiang Province, China. The tall, thin-walled species possessed unique features including a combination of robust longitudinal monaxons, fine sub-transverse monaxons, and minute hexactine and hetera...
Botting, J.P., Muir, L.A., Xiao, S., Li, X. & Lin, J.-P. 2012: Evidence for spicule homology in calcareous and siliceous sponges: biminerallic spicules in Lenica sp. from the Early Cambrian of South China. Lethaia, Vol. 45, pp. 463–475.
The relationships of the extant sponge classes, and the nature of the last common ancestor of all sponges, are cu...
The monotypic Burgess Shale sponge genus Takakkawia Walcott, 1920, has been previously assigned to the Protomonaxonida, despite showing several unique features. A reassessment of the genus, including restudy of previously described material and the addition of new material that includes partially disarticulated specimens, has allowed a greatly modi...
A new odontopleurid trilobite, Meadowtownella serrata, is described from a new Llanvirnian (Darriwilian; Didymograptus murchisoni Biozone) locality in the Builth Inlier, Mid-Wales. This unusually spinose species displays a remarkable array of pygidial spines and extends the stratigraphic range of the genus back into the Abereiddian. Benthic faunas...
Palaeoscolecidan worms are rare, Early Palaeozoic fossils with uncertain affinities within the Ecdysozoa. They are locally abundant in the Cambrian and scattered in the Early Ordovician, but very sparse thereafter. Forty‐four specimens have been collected from the Middle Ordovician of the Builth‐Llandrindod Inlier of Mid Wales and include well‐pres...
A new sponge fauna has been discovered in silty mudstone of the early Migneintian (late Tremadocian, Ordovician) of North Wales. The assemblage is dominated by reticulosan hexactinellids, including several species bearing parietal gaps; this feature is common among Cambrian hexactinellids, but is rare in Ordovician faunas. Of particular significanc...
Six specimens of a strongly curved, cylindrical hexactinellid sponge have been recovered from the TommotianAtdabanian Hetang Biota of South China, and are described as Decumbispongia yuani gen. et sp. nov. The robust, thick-walled sponge shows no evidence of an osculum or basal structures, and the body form is inconsistent with an upright, filter-f...