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A new species of bubble-headed trilobite, Staurocephalus oliveae sp. nov., is described from 250 specimens collected from a recently discovered locality in the Sholeshook Limestone Formation (upper Katian, Upper Ordovician; Ashgill, Cautleyan Zone 1 equivalent) in the village of Llanddowror, Carmarthenshire. Although no fully articulated specimens were found, the material is the most abundant and well-preserved of any Ordovician Staurocephalus species known from Britain and Ireland, and enables the complicated 3D structure of its cephalon to be reconstructed. Staurocephalus oliveae sp. nov. is the earliest representative of the genus recorded in Avalonia, following its presumed migration into the area from Laurentia or Baltica. Recognition of this Cautleyan species resurrects the potential usefulness of the long established, widely distributed, Staurocephalus clavifrons as a latest Katian (= Rawtheyan) indicator.

Morphological disparity and taxonomic richness are two major aspects of evolution used to understand biodiversity changes. These metrics are often decoupled in time, particularly during the early history of clades. To assess the pervasiveness of this pattern during the post-acme of a clade, both morphological disparity and taxonomic diversity of the well-documented Devonian trilobites from North Africa were analysed. Morphospace occupancy and body size were estimated and compared to genus richness through time. This study highlights that, during the Early Devonian, morphological disparity of the pygidium and the cephalon strongly increased, whereas cranidium disparity remained low. Interestingly, the pygidium and cephalon morphological dynamics were decoupled. Taxonomic diversity also increased. Then, the Middle Devonian anoxic events affected the trilobite communities with simultaneous drastic loss of both morphologies and taxonomic richness. This coupling between diversity and disparity dynamics could be explained by either the intensity of extinctions or strong internal constraints. Finally, the weak Famennian recovery of both disparity and diversity did not reach the Early Devonian levels, thus making trilobites a 'dead clade walking' during their late evolutionary history. Devonian trilobite families are thus identified as 'Bottom-Heavy Clades', characterised by a diversity peak at the end of their history. [ABSTRACT FROM AUTHOR]

Floian, Early Ordovician trilobites are systematically described and revised based on new material from the middle part of the Duoquanshan Formation of the Shihuigou area, northern Qinghai Province, northwest China. The fauna that lived on the shallow-water carbonate platform comprises three species belonging to two families, i.e., Tsaidamaspis diarmatus, Zhiyia tsinghaiensis, and Liexiaspis sp. indeterminate. It exhibits a strong endemicity to the Olongbluk terrane. The new isoteline genus Zhiyia is established on the basis of the material from the Olongbluk terrane and South China palaeoplate, and is characterized by its: (i) almost obsolete cephalic and pygidial axial furrows; (ii) flattened anterior border and narrow (sag., exsag.) occipital ring; (iii) bilobed hypostome with a shallow median notch and a small triangular median projection; (iv) subsemicircular pygidium with wide pygidial axis and border. Faunal evidence indicates that the palaeogeographic position of the Olongbluk terrane may have been situated closer to the South China palaeoplate rather than the North China palaeoplate during the Floian.

Species of the odontopleurine trilobites Leonaspis Richter & Richter, 1917 and Kettneraspis Prantl & Přibyl, 1949 from the Silurian and the Devonian have been studied. Dorsal tubercle and spine patterns are highly variable and deemed of restricted use. The pygidial segmentation of Leonaspis is consistent among species in the development of a rudimentary inner pleural ridge or wide space between the pleural ridge and the axis. In Kettneraspis, an inner pleural ridge is never developed but the primitive condition is to have a space here. The formation of this ridge and an auxiliary interior border spine pair was arrested prematurely during ontogeny in species assigned to Kettneraspis at present. The type species of the monotypic Rupbachella Alberti, 2021 is regarded as a juvenile Leonaspis. Bruthansovaspis gen. nov. (type species: Acidaspis roemeri Barrande, 1852) is proposed for a Silurian clade of former Kettneraspis species from peri-Gondwana, that are principally distinct in lacking the combination of stunted anterior and macropleural posterior, thoracic segments synapomorphic of Kettneraspis and Leonaspis. The types of Bruthansovaspis roemeri, Bruthansovaspis dormitzeri (Hawle & Corda, 1847), Bruthansovaspis dumortieri (Hawle & Corda, 1847), Kettneraspis propinqua (Barrande, 1852), Kettneraspis lindackeri (Hawle & Corda, 1847) and Bruthansovaspis zenkeri (Hawle & Corda, 1847) (= B. dormitzeri) from the Silurian of the Barrandian, all of which had previously been lumped as dormitzeri, are figured and their classifications revisited. Exceptionally well-preserved odontopleurine material from the Devonian in the Tafilalt (southern Morocco) is described as Kettneraspis freitagi sp. nov. Leonaspis' strix Lütke, 1965 from the Devonian of the Harz Mountains is in need of revision; a previously tentatively assigned pygidium does not belong in that genus. [ABSTRACT FROM AUTHOR]

A new species of bubble-headed trilobite, Staurocephalus oliveae sp. nov., is described from 250 specimens collected from a recently discovered locality in the Sholeshook Limestone Formation (upper Katian, Upper Ordovician; Ashgill, Cautleyan Zone 1 equivalent) in the village of Llanddowror, Carmarthenshire. Although no fully articulated specimens were found, the material is the most abundant and well-preserved of any Ordovician Staurocephalus species known from Britain and Ireland, and enables the complicated 3D structure of its cephalon to be reconstructed. Staurocephalus oliveae sp. nov. is the earliest representative of the genus recorded in Avalonia, following its presumed migration into the area from Laurentia or Baltica. Recognition of this Cautleyan species resurrects the potential usefulness of the long established, widely distributed, Staurocephalus clavifrons as a latest Katian (= Rawtheyan) indicator. [ABSTRACT FROM AUTHOR]

The ability to enrol for protection is an effective defensive strategy that has convergently evolved multiple times in disparate animal groups ranging from euarthropods to mammals. Enrolment is a staple habit of trilobites, and their biomineralized dorsal exoskeleton offered a versatile substrate for the evolution of interlocking devices. However, it is unknown whether trilobites also featured ventral adaptations for enrolment. Here, we report ventral exoskeletal adaptations that facilitate enrolment in exceptionally preserved trilobites from the Middle Ordovician Walcott–Rust Quarry in New York State, USA. Walcott–Rust trilobites reveal the intricate three-dimensional organization of the non-biomineralized ventral anatomy preserved as calcite casts, including the spatial relationship between the articulated sternites (i.e. ventral exoskeletal plates) and the wedge-shaped protopodites. Enrolment in trilobites is achieved by ventrally dipping the anterior margin of the sternites during trunk flexure, facilitated by the presence of flexible membranes, and with the close coupling of the wedge-shaped protopodites. Comparisons with the ventral morphology of extant glomerid millipedes and terrestrial isopods reveal similar mechanisms used for enrolment. The wedge-shaped protopodites of trilobites closely resemble the gnathobasic coxa/protopodite of extant horseshoe crabs. We propose that the trilobites' wedge-shaped protopodite simultaneously facilitated tight enrolment and gnathobasic feeding with the trunk appendages. [ABSTRACT FROM AUTHOR]

A poorly diverse trilobite assemblage is described from the Tremadocian strata (Tangissart Member of the Mousty Formation; Chevlipont Formation) of the Brabant Massif. These specimens represent so far the oldest trilobite record from Belgium. The recorded taxa, identified as Platypeltoides cf. croftii , Macropyge ? sp., and Asaphidae indet., resemble contemporaneous trilobites from other parts of Avalonia (e.g., Wales, Shropshire) as well as some others from high-latitude Gondwana (Morocco). The nileid species P. cf. croftii suggests that the Tangissart Member of the Mousty Formation was likely deposited in deeper offshore environments on the open shelf of Avalonia. Morphological similarities between P. cf. croftii from the Brabant Massif, P. croftii from the UK, and P. magrebiensis from Morocco suggest close faunal interchange across the narrow Rheic Ocean. [ABSTRACT FROM AUTHOR]

The Lower Ordovician Fezouata Shale is renowned for its exceptionally-preserved euarthropod fossils including numerous species of trilobites, some of which show remains of appendages and traces of the digestive system. Herein, we describe the early developmental stages of at least nine trilobite species from the Tremadocian strata of the Fezouata Shale, namely Platypeltoides magrebiensis , Nileus deynouxi , Symphysurus ebbestadi , Asaphellus sp., Megistaspis (Ekeraspis) hammondi , Orometopus sp., Anacheirurus adserai , Bavarilla zemmourensis , Indiligens sp., and several specimens of undetermined protaspides. This study considerably expands our knowledge of the development of early Ordovician trilobites. The preservation of appendages in the early stages of N. deynouxi and S. ebbestadi , and remains of the digestive tract in the latter species, suggests that some immature trilobites had similar morphology and anatomy as the adult individuals. Early developmental stages of Indiligens sp. might have fed and/or hidden on graptolites and demosponges. The extraordinarily large size of the early post-embryonic stages of P. magrebiensis , S. ebbestadi , Orometopus sp., Asaphellus sp., and undetermined protaspides suggests that these trilobites might have hatched from yolk-rich eggs. The presence of several trilobite species with notably large post-embryonic stages in the Fezouata Shale might be explained by seasonal or low productivity in the high-latitude margin of Gondwana. [ABSTRACT FROM AUTHOR]

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Eccaparadoxides is a geographically widely distributed trilobite genus that occurs in the middle part of the Cambrian System. However, the systematically important morphologic characteristics that can be used to differentiate taxa are often problematical in their application. A review of the large number (over 30) of significant species or forms assigned to Eccaparadoxides clearly indicates that only the pygidia offer fairly reliable morphologic criteria that can be used taxonomically and phylogenetically. The pygidia allow for recognition of four different morphological groups (pusillus, lamellatus, pradoanus and asturianus) of which the asturianus group can only be questionably assigned to the genus. Species known only from cranidia cannot be assigned to Eccaparadoxides with certainty. This study refines the biostratigraphy for the interval from the upper Wuliuan to the middle Drumian and shows that this interval brackets the range of most Eccaparadoxides species. The genera or subgenera Baltoparadoxides, Rejkocephalus and Macrocerca are evaluated. Eccaparadoxides zelus, E. epimetheus and Eccaparadoxides? hestia are newly proposed species. [ABSTRACT FROM AUTHOR]

Tuff‐bearing upper Cambrian to lowermost Ordovician strata on Ko Tarutao island, Satun province, southernmost peninsular Thailand, contain a rich trilobite fauna relevant to global biostratigraphy, peri‐Gondwanan palaeogeography and shifting evolutionary mode. This area of Sibumasu, a lower Palaeozoic marginal Gondwanan terrane, is shown to have been closely associated with Australia, North China (Sino‐Korea) and other continental fragments from the supercontinent's northern equatorial sector, including South China at that time. Shared faunas also suggest a Kazakhstani and Laurentian association. Collections from eight sections yielded 10 newly discovered species and one new genus from ancient shoreface and inner shelf siliciclastic deposits. With the new taxa and revision of taxa known previously, we refine the age of the upper two formations of the Tarutao Group to the middle of Cambrian Stage 10, and lower–middle Tremadocian. Two biozones are erected for Sibumasu: the Eosaukia buravasi Zone, encompassing all Cambrian sections from Ko Tarutao, and the Asaphellus charoenmiti Zone, encompassing the Tremadocian fauna discussed herein. The new genus is Tarutaoia and new species are Tsinania sirindhornae, Pseudokoldinioidia maneekuti, Pagodia? uhleini, Asaphellus charoenmiti, Tarutaoia techawani, Jiia talowaois, Caznaia imsamuti, Anderssonella undulata, Lophosaukia nuchanongi and Corbinia perforata. Other taxa reported for the first time from Tarutao are Mansuyia? sp., Parakoldinioidia callosa Qian, Pseudagnostus sp., Homagnostus sp., Haniwa mucronata Shergold, Haniwa sosanensis? Kobayashi, Lichengia simplex Shergold, Pacootasaukia sp., Wuhuia? sp., Plethopeltella sp., Apatokephalus sp., Akoldinioidia sp. 1 and Koldinioidia sp. [ABSTRACT FROM AUTHOR]

Three shelf-derived boulders in debris flow conglomerates of the Downes Point Member of the Shallow Bay Formation of western Newfoundland yielded the first record of an Upper Cambrian shelf-margin trilobite fauna from the Elvinia Zone (Steptoean; Jiangshanian) in eastern North America. The fauna is dominated by the "catillicephalid" Buttsia Wilson, 1951, and resembles trilobite biofacies from microbial buildups in the Gatesburg Formation of Pennsylvania. It is correlative with the Cliffia latagenae Subzone, which is the youngest Steptoean biostratigraphic unit in the Gatesburg. New taxa are Triorygma burkhalteri gen. et sp. nov. and Buttsia trema sp. nov. [ABSTRACT FROM AUTHOR]

Efficient extraction of oxygen from ambient waters played a critical role in the development of early arthropods. Maximizing gill surface area enhanced oxygen uptake ability but, with gills necessarily exposed to the external environment, also presented the issue of gill contamination. Here we document setae inserted on the dorsal surface of walking legs of the benthic-dwelling middle Cambrian Olenoides serratus and on the gill shaft of the Late Ordovician Triarthrus eatoni. Based on their physical positions relative to gill filaments, we interpret these setae to have been used to groom the gills, removing particles trapped among the filaments. The coordination between setae and gill filaments is comparable to that seen among modern crustaceans, which use a diverse set of setae-bearing appendages to penetrate between gill filaments when grooming. Grooming is known relatively early in trilobite evolutionary history and would have enhanced gill efficiency by maximizing the surface area for oxygen uptake. [ABSTRACT FROM AUTHOR]

The new scutelluid trilobite genus Mitroplax, with type species M. enormis (Etheridge, 1894), occurs in Pragian to early Emsian strata at several localities in Victoria, Australia. The genus is characterized by a remarkably long and flat anterior cephalic border, a glabella that expands almost uniformly forward from the occipital furrow, a subcircular S2 discrete from S1, and a pygidium with the well-rounded paired pleural ribs and the posterior bifurcations of the postaxial rib tapering almost to a point distally. The type species grew to exceptional size, as indicated by the dimensions of the largest pygidia. Another scutelluid from the Lochkovian of Victoria, ‘Goldius’ greenii Chapman, 1914, is revised and assigned to Spiniscutellum Šnajdr, 1960. This species and a related one from the Lochkovian of the Bungonia district of south-eastern New South Wales are the first records of Spiniscutellum from eastern Gondwana. Spiniscutellum (Lochkovopeltis) Hörbinger, 2004 is regarded as a junior synonym of Spiniscutellum. [ABSTRACT FROM AUTHOR]

Species of Catillicephala are known from sites around the mid-Cambrian margin of Laurentian North America, including Vermont, Quebec, Newfoundland and North Greenland. Catillicephala cifellii sp. nov. is from the Downes Point Member of the Shallow Bay Formation (Cow Head Group) in western Newfoundland. It occurs in three shelf margin-derived boulders in debris flow conglomerates that accumulated in a continental slope setting. The associated trilobites and agnostoid arthropods, including Ptychagnostus aculeatus and Megagnostus glandiformis, indicate a correlation with the Lejopyge laevigata Zone. As such, C. cifellii is among the oldest representatives of the genus, and is early Guzhangian in age.

Arthropods are characterized by having an exoskeleton, paired jointed appendages and segmented body. The number and shape of those segments vary dramatically and unravelling the evolution of segmentation is fundamental to our understanding of arthropod diversification. Because trilobites added segments to the body post-hatching which were expressed and preserved in biomineralized exoskeletal sclerites, their fossil record provides an excellent system for understanding the early evolution of segmentation in arthropods. Over the last 200 years, palaeontologists have hypothesized trends in segment number and allocation in the trilobite body, but they have never been rigorously tested. We tabulated the number of segments in the post-cephalic body for over 1500 species, selected to maximize taxonomic, geographical and temporal representation. Analysis reveals long-term shifts in segment number and allocation over the 250-million-year evolutionary history of the clade. For most of the Palaeozoic, the median number of segments in the body did not change. Instead, the total range decreased over time and there was long-term increase in the proportion of segments allocated to the fused terminal sclerite relative to the articulated thoracic region. There was also increased conservation of thoracic segment number within families. Neither taxonomic turnover nor trends in functionally relevant defensive behaviour sufficiently explain these patterns. [ABSTRACT FROM AUTHOR]

The deep-water Aulacopleura koninckii Assemblage in the lower Homerian (T. testis Sub-Biozone) "Aulacopleura shales" strata at the classical 'Barrande's pits' locality on the Černidla hillside at Loděnice in the Prague Basin is supplemented by the addition of two new trilobite taxa, viz. Exallaspis? perunicana sp. n. and Kosovoproetus? aff. praecursor (Přibyl & Vaněk 1987). The palaeogeographic distribution of Exallapis Ramsköld & Chatterton, 1991 is extended by the occurrence of Exallaspis? perunicana sp. n. and the younger Exallaspis sp. in the upper Homerian (the P. parvus – G. nassa Biozone) of the Prague Basin, reflecting the faunal migration between the southern shelf of the Baltica palaeocontinent and the Perunica microcontinent accross the Rheic Ocean. The small dimensions of the K.? aff. praecursor exoskeletons compared to those of K.? praecursor in the bordering shallow-water Liolalax–Sphaerexochus– Cheirurus Assemblage represent another example of adaptive nanism in trilobites of the Aulacopleura-Raphiophorus Biofacies. [ABSTRACT FROM AUTHOR]

Pilekiines are the earliest diverging members of the successful trilobite family Cheiruridae. The pilekiine genus Anacheirurus is characterized by sub-quadratic to sub-oval glabella, pitted genae, and a distinct trunk with elongated pleural spines in its posterior part. Anacheirurus adserai is a common component of the Fezouata Shale (Lower Ordovician, Morocco), where it was intially included into several species of the genus Lehua. This assignment and taxonomic over-splitting created confusion, overestimated cheirurid diversity at this locality, and simultaneously underestimated morphological variability within A. adserai. In this contribution we examine new material of A. adserai from the Fezouata Shale, clarifying its morphology and systematics. A detailed re-description of the species shows that Anacheirurus is distinct from Lehua, the latter being a more derived member of Cheiruridae. The comparison of Anacheirurus with other pilekiines shows that morphological variability within this subfamily is mostly constrained to the trunk region. Exceptionally preserved specimens of A. adserai from the Fezouata Shale show details of appendages, revealing the endopodite and exopodite morphologies in early members of Cheiruridae. The endopodite of A. adserai is unique among trilobites in possessing comparatively longer distal podomeres 5 and 6, but otherwise, it has the same general morphology as other described trilobite endopodites. The exopodite morphology of A. adserai shows characters typical of some Cambrian species but differs in several aspects from those known in post-Cambrian taxa. It is concluded that trilobite exopodite morphology was probably more variable than the endopodite morphology, which remains rather conservative across different taxa. Morphological diversity of trilobite exopodites in post-Cambrian taxa might be related to ecological escalations during the Ordovician biodiversification and the transition between Cambrian and Ordovician trilobite faunas.

Thirteen species of trilobites are described from two different faunas in late Katian and Rhuddanian strata respectively of the Arndell Sandstone in the Florentine Valley of southern Tasmania. The late Katian fauna includes the trinucleid Auritolithus corbetti gen. et sp. nov., the calymenids Salterocoryphe? bailliei sp. nov. and Vietnamia hyron sp. nov., and limited material of the lichid Amphilichas? and two indeterminate asaphids. Auritolithus, characterized by the markedly subpentagonal outline of the cephalon, cannot be assigned to any of the existing trinucleid subfamilies but seems closely related to forms occurring in the Upper Ordovician of the Precordillera Basin of Argentina. Salterocoryphe? bailliei, the first record of a member of the Colpocoryphinae from Australia, is considerably younger stratigraphically than other occurrences of that genus in the middle to upper Darriwilian of France and the Iberian Peninsula. The Rhuddanian fauna from the Arndell Sandstone includes the illaenid Pepodes agrestis gen. et sp. nov., the scutelluid Japonoscutellum senectum sp. nov., the brachymetopid Niuchangella agastor sp. nov., the encrinurids Arndellaspis oryxis gen. et sp. nov. and Cromus?, the calymenid Gravicalymene clarkei sp. nov. and the homalonotid Brongniartella calveri sp. nov. Niuchangella is accepted as a senior synonym of Radnoria. Arndellaspis is not closely related to other members of the Encrinurinae from Australia but shares unique derived characters of the pygidium with Encrinurus (s.s.) and Wallacia, occurring in Laurentia, Avalonia and Baltica, and Arndellaspis may share common ancestry with Encrinurus. The commonly broadly interpreted Gravicalymene is revised and restricted to a smaller number of closely related species from the Darriwilian to middle Telychian. The faunal affinities of the trilobites suggest links with terranes that lay along the north-western margin of Gondwana and with South America in the Late Ordovician, and with South China and possibly Laurentia in the early Silurian. [ABSTRACT FROM AUTHOR]

Some exceptional paleontological trilobite sites located in the Sierra Norte de Sevilla UNESCO Global Geopark are presented herein, together with an analysis of their geotourism / geotrail potential and a proposal for geoconservation. The sites are of Marianian age, a regional stage and age of the Cambrian Mediterranean Subprovince which was defined within the territory of the Geopark. This area constitutes the type area of six trilobite genera and twelve trilobite species and other associated faunas (hyolith, serpulid and echinoderm species).

The Suborder Cheirurina is a diverse trilobite clade, ranging from the latest Cambrian to the Devonian. The Ordovician paleobiogeography of the group is explored based on the distribution of genera. Major tectonostratigraphic units and Ordovician ages (Tremadocian, Floian, Dapingian, Darriwilian, Sandbian and Katian) were recorded for each genus. Cluster analysis (agglomerative hierarchical clustering) and non-metric multidimensional scaling (NMDS) were performed independently for each Ordovician age. The results characterize the Ordovician distribution of Cheirurina as two main general patterns. First, there was a marked decrease in endemicity through the Ordovician. Second, latitude seems to play a significant role in the distribution of Cheirurina, at both genus and family levels. Pliomerids and encrinurids occur overwhelmingly at low latitudes. The two main exceptions are Placoparia (Pliomeridae) and Dindymene (Encrinuridae), which are distributed in high latitude terranes. Within Cheiruridae, members of Eccoptochilinae are typical of high latitude Gondwana. Conversely, acanthoparyphines, cheirurines, deiphonines, heliomerines and sphaerexochines are mainly distributed in low or intermediate latitudes. • The distribution of the largest Ordovician trilobite clade is explored in this study. • Four major faunal provinces are recognized throughout the Ordovician. • The distribution of the group is highly influenced by the latitude. • A reduction of provincialism is observed through the Ordovician. • During the Katian the group showed a more cosmopolitan distribution than before. [ABSTRACT FROM AUTHOR]

Serrodiscus is considered one of the first trilobite genera showing global distribution during the Cambrian and a key component for the international correlation of this period. However, this genus encompasses many poorly documented species, ambiguous diagnoses or characters that depend on taphonomy and deformation based on one single specimen. Thus, in the present work, we carry out a systematic overview of the genus, evaluating the taphonomic and deformational variability of a large number of specimens from the same locality and horizon. In addition, and due to the difficulty of making synonymy decisions, closely related species are grouped into three different groups taking into consideration morphologic, stratigraphic and palaeobiogeographic data: bellimarginatus group, scattered along Avalonian and western Gondwanan margins; speciosus group, extending over the Laurentian domain; and daedalus group, being restricted to eastern Gondwana (Australia), North China and, partially, Siberia. Besides, the regional correlation through the early Cambrian of Iberia is arranged, describing specimens from both Ossa-Morena and Central Iberian zones, with a time interval restricted to the upper Marianian Stage. Regarding the biostratigraphy and palaeobiogeography of the genus, Serrodiscus has been reported from Laurentia, Taconic Allochthon, Greenland, Baltica, western and eastern Avalonia, western Gondwana margin, Siberia, North China and eastern Gondwana, with a temporal distribution along with the Cambrian Series 2. [ABSTRACT FROM AUTHOR]

Middle to Late Ordovician (late Darriwilian –early Katian) trilobites from trinucleid, raphiophorid and cyclopygid biofacies are for the first time documented from the West Balkhash and North Betpak‐Dala regions in south‐central Kazakhstan. Twenty taxa are described; six of them, that is, Miaopopsis sokyrensis sp. nov., Pricyclopyge keralensis sp. nov., Kongqiaoheia sarytumensis sp. nov., Shumardia karasaiensis sp. nov., Triarthrus akkermensis sp. nov. and Mynaralaspis perforata gen. et sp. nov. are new to science. Six taxa are reported in open nomenclature and two are not identified on a genus level. A new trinucleid Subfamily Nanshanaspinae is designated. Newly discovered faunas show strong links to contemporaneous faunas of Tarim and, to a lesser degree, to those of South China and the Australian sector of Gondwana. Based on results of cluster analysis (Dice similarity), trilobite associations of raphiophorid biofacies from the Chu‐Ili and Stepnyak terranes form a single first‐order cluster with little similarity to the trilobite associations characteristic of the nielid and illenid–cheirurid biofacies from the Stepnyak and Tarbagatai regions. Also, there is no distinct interaction between the pelagic Degamella–Pricyclopyge association and other trilobite faunas. The geographical distribution of raphiophorid and trinucleid trilobites described in this paper, including species of genera Malongullia?, Miaopopsis, Nanshanaspis and Kongqiaoheia, supports Ordovician palaeogeographical reconstructions showing the Kazakh terranes as part of a huge archipelago situated on both sides of the equator in proximity to the west coast of Gondwana and positioned a considerable distance from the Siberia continent and its associated terranes. [ABSTRACT FROM AUTHOR]

Hypermecaspidinae (Trilobita; Olenidae) constitutes a monophyletic clade but its internal relationships are unknown and its phylogenetic position with respect to the rest of Olenida has recently been questioned. In order to propose a phylogenetic hypothesis, and to resolve the status of the group, a cladistic analysis of the hypermecaspidinids was performed. One hundred characters (28 quantitative and 72 qualitative) of holaspid morphology were coded for 30 taxa (18 Hypermecaspidinae). Two groups of analyses were performed considering the percentage of missing data. For each group of analyses, discrete partition was treated as totally unordered or as with some characters (describing intensity, position or shape of certain structures) ordered. The analyses were performed with TNT using maximum parsimony and implied weighting as optimality criteria. Hypermecaspididae constitutes a monophyletic family within Olenida, and is the sister group of pterocephaliids. In light of the present analysis, an emended diagnosis of the family is provided. The results suggest that 'Tropidopyge' broeggeri should be reassigned to Hypermecaspis. Hypermecaspidids would have originated in the late Furongian and rapidly diversified across different parts of the world in the Early Ordovician. With the exception of H. armata and H. chamarrensis, the species of Hypermecaspis have a proportionately large pygidium with a wide doublure and a distinct postaxial field. An evolutionary trend toward an increase in the size of these characters is observed in the genus. The cranidium shows a tendency toward enlargement of the glabella and the palpebral lobes, together with a narrowing of the interocular genae. [ABSTRACT FROM AUTHOR]

Fantasticolithus isabelae, a new genus and species of trinucleid trilobite from the Lower Ordovician of Peru, developed anterior ventral cephalic 'props' that are unique in the Trilobita. Its broad and finely pitted genal prolongations are unlike those of any previously known trinucleid, and closely resemble those invariably present in the unrelated Family Harpetidae, supporting the hypothesis that the two families shared a similar mode of life, as 'trinucleimorphs'. Fantasticolithus was probably derived from an early trinucleid with a marginal girder, although it resembles younger genera such as Marrolithus in its enlarged anterolateral pits. The new genus provides a natural experiment to test the hypothesis of trinucleimorphs as filter feeders that used a ventral feeding chamber. The role of the pitted marginal fringe as an egress for feeding currents is discussed. Fantasticolithus had an unusual anterior vent that provided a simpler exit mechanism for feeding currents than the pitted fringe. However, the pits were probably important for trinucleids that lacked an anterior arch. [ABSTRACT FROM AUTHOR]

Trilobites achieved their maximum generic diversity in the Cambrian, but the peak of morphological disparity of their cranidia occurred in the Middle to Late Ordovician. Early to middle Cambrian trilobites with a specialized cephalon are rare, especially among the ptychoparioids, a group of libristomates featuring the so-called “generalized” bauplan. Here we describe an unusual ptychopariid trilobite Phantaspis auritus gen. et sp. nov. from the middle Cambrian (Miaolingian, Wuliuan) Mantou Formation in the Shandong Province, North China. This new taxon is characterized by a cephalon with an extended anterior area of double-lobate shape resembling a pair of rabbit ears in later ontogenetic stages; a unique type of cephalic specialization that has not been reported from other trilobites. Such a peculiar cephalon as in Phantaspis provides new insights into the variations of cephalic morphology in middle Cambrian trilobites, and may represent a heuristic example of ecological specialization to predation or an improved discoidal enrollment.

Species of Catillicephala are known from sites around the mid-Cambrian margin of Laurentian North America, including Vermont, Quebec, Newfoundland and North Greenland. Catillicephala cifellii sp. nov. is from the Downes Point Member of the Shallow Bay Formation (Cow Head Group) in western Newfoundland. It occurs in three shelf margin-derived boulders in debris flow conglomerates that accumulated in a continental slope setting. The associated trilobites and agnostoid arthropods, including Ptychagnostus aculeatus and Megagnostus glandiformis, indicate a correlation with the Lejopyge laevigata Zone. As such, C. cifellii is among the oldest representatives of the genus, and is early Guzhangian in age. [ABSTRACT FROM AUTHOR]

122 specimens of the trilobite species Cyrtoproetus (Cyrtoproetus) moravicus (PŘIBYL, 1950), whose fossils come from the culm of Drahany Upland, are deposited in the paleontological collection of the Regional Museum in Olomouc. Its systematic affi liation, occurrences within the culm of the Drahany Upland and biostratigraphic range are discussed. [ABSTRACT FROM AUTHOR]

The first Silurian trilobite known with soft parts preserved, a Dalmanites species, is described from the Herefordshire Lagerstätte. Biramous appendages and much of the alimentary system are evident. High‐fidelity three‐dimensional preservation reveals a novel arrangement of the exopod, in which successive filaments are connected by a presumed membrane. This morphology explains a misinterpretation of the exopod as supporting spiral structures, originally reported nearly 150 years ago. Comparison with other trilobite limbs indicates that the exopod morphology of Dalmanites is present in other members of Phacopida. The function of the exopod is considered to be primarily respiratory. [ABSTRACT FROM AUTHOR]

Pilekiines are the earliest diverging members of the successful trilobite family Cheiruridae. The pilekiine genus Anacheirurus is characterized by sub-quadratic to sub-oval glabella, pitted genae, and a distinct trunk with elongated pleural spines in its posterior part. Anacheirurus adserai is a common component of the Fezouata Shale (Lower Ordovician, Morocco), where it was intially included into several species of the genus Lehua. This assignment and taxonomic over-splitting created confusion, overestimated cheirurid diversity at this locality, and simultaneously underestimated morphological variability within A. adserai. In this contribution we examine new material of A. adserai from the Fezouata Shale, clarifying its morphology and systematics. A detailed re-description of the species shows that Anacheirurus is distinct from Lehua, the latter being a more derived member of Cheiruridae. The comparison of Anacheirurus with other pilekiines shows that morphological variability within this subfamily is mostly constrained to the trunk region. Exceptionally preserved specimens of A. adserai from the Fezouata Shale show details of appendages, revealing the endopodite and exopodite morphologies in early members of Cheiruridae. The endopodite of A. adserai is unique among trilobites in possessing comparatively longer distal podomeres 5 and 6, but otherwise, it has the same general morphology as other described trilobite endopodites. The exopodite morphology of A. adserai shows characters typical of some Cambrian species but differs in several aspects from those known in post-Cambrian taxa. It is concluded that trilobite exopodite morphology was probably more variable than the endopodite morphology, which remains rather conservative across different taxa. Morphological diversity of trilobite exopodites in post-Cambrian taxa might be related to ecological escalations during the Ordovician biodiversification and the transition between Cambrian and Ordovician trilobite faunas. [ABSTRACT FROM AUTHOR]

The morphology and ontogeny of the trilobite Changaspis elongata based on 216 specimens collected from the Lazizhai section of the Balang Formation (Stage 4, Series 2 of the Cambrian) in Guizhou Province, South China are described. The relatively continuous ontogenetic series reveals morphological changes, and shows that the species has seventeen thoracic segments in the holaspid period, instead of the sixteen as previously suggested. The development of the pygidial segments shows that their number gradually decreases during ontogeny. A new dataset of well-preserved specimens offers a unique opportunity to investigate developmental traits after segment addition is completed. The ontogenetic size progressions for the lengths of cephalon and trunk show overall compliance with Dyar’s rule. As a result of different average growth rates for the lengths of cephalon, trunk and pygidium, the length of the thorax relative to the body shows a gradually increasing trend; however, the cephalon and pygidium follow the opposite trend. Morphometric analysis across fourteen post-embryonic stages reveals growth gradients with increasing values for each thoracic segment from anterior to posterior. The reconstruction of the development traits shows visualization of the changes in relative growth and segmentation for the different body parts. The new dataset and growth gradient of the trunk suggest that the thoracic segment growth dynamics of early Cambrian to Silurian trilobites follow the same general continuous, steady-state growth gradient decreasing from posterior to anterior.

The lower Cambrian (Series 2) White Point Conglomerate (WPC) on Kangaroo Island, South Australia contains exotic clasts representing a diverse array of lithologies, including metamorphics, chert, sandstone, and abundant carbonates, notably archaeocyath-rich bioclastic limestone. Acetic acid digestion of the WPC bioclastic limestone clasts reveals a diverse shelly fauna. This assemblage includes abundant organophosphatic brachiopods such as Cordatia erinae Brock and Claybourn gen. et sp. nov., Curdus pararaensis, Eodicellomus elkaniformiis, Eohadrotreta sp. cf. E. zhenbaensis, Eoobolus sp., Kyrshabaktella davidii, and Schizopholis yorkensis. Additional shelly taxa include the solenopleurid trilobite Trachoparia? sp., the tommotiids Dailyatia odyssei, Dailyatia decobruta Betts sp. nov., Kelanella sp., and Lapworthella fasciculata, spines of the bradoriid arthropod Mongolitubulus squamifer, and several problematica, such as Stoibostrombus crenulatus and a variety of tubular forms. The upper age limit for the WPC is constrained by biostratigraphic data from the overlying Marsden Sandstone and Emu Bay Shale, which are no younger than the Pararaia janeae Trilobite Zone (Cambrian Series 2, Stage 4). The shelly fossil assemblage from the WPC limestone clasts indicates an upper Dailyatia odyssei Zone (= Pararaia tatei to lower P. janeae trilobite zones), equivalent to the Atdabanian–early Botoman of the Siberian scheme. This contrasts with the previously suggested late Botoman age for the limestone clasts, based on the diverse archaeocyath assemblage. The minor age difference between the WPC and its fossiliferous limestone clasts suggests relatively rapid reworking of biohermal buildups during tectonically-active phases of deposition in the Stansbury Basin.

Euarthropods have a tough exoskeleton that provides crucial protection from predation and parasitism. However, this is restrictive to growth and must be periodically moulted. The moulting sequence is well-known from extant arthropods, consisting of: (i) the long inter-moult stage, in which no changes occur to the hardened exoskeleton; (ii) the pre-moult stage where the old exoskeleton is detached and the new one secreted; (iii) exuviation, when the old exoskeleton is moulted; and (iv) the post-moult stage during which the new exoskeleton starts as soft, thin, and partially compressed and gradually hardens to the robust exoskeleton of the inter-moult stage. Trilobite fossils typically consist of inter-moult carcasses or moulted exuviae, but specimens preserving the post-moult stage are rare. Here we describe nine specimens assigned to Symphysurus ebbestadi representing the first group of contemporaneous fossils collected that preserve all key stages of the moulting process in one taxon, including the post-moult stage. They were collected from a single lens in the Tremadocian part of the Fezouata Shale Formation, Morocco. Based on cephalic displacement and comparison to other trilobite moults, one specimen appears to represent a moulted exoskeleton. Four specimens are typical inter-moult carcasses. Four others are wrinkled and flattened, with thin exoskeletons compared to inter-moult specimens, and are considered post-moult individuals. These S. ebbestadi specimens illuminate the preservation and morphology of the post-moulting stage, characterised by strong anterior-posterior exoskeleton wrinkling, as well as overall body flattening and reduced visibility of thoracic articulations. Being found in the same lens, these specimens likely represent the first preserved in-the-act mass moulting event. The displayed sequence of moulting suggests the moulting process in trilobites was comparable to modern arthropods, and conserved within euarthropod evolutionary history.

A new cornuproetine trilobite with nine thorax segments from upper Emsian strata in the NE Draa Valley (southern Morocco) is described as Timsaloproetus alissae sp. nov. Morphological evidence is reviewed to suggest a semi-endobenthic life mode, possibly with a carnivorous feeding habit. [ABSTRACT FROM AUTHOR]

Trilobites offer almost unparalleled insight into the growth and development of fossil ecdysozoans. Here I use newly collected material of Elrathia kingii (Meek) to estimate growth rates and describe shape change over the ontogeny of E. kingii. Well‐preserved, articulated specimens from all post‐embryonic stages were collected from a 1.5 m interval of the upper Wheeler Formation (Miaolingian Series, Cambrian) in western Utah (USA), and size and landmark‐based shape data were digitized from photographs. Growth rates were estimated and compared with previously published data on the Silurian trilobite, Aulacopleura koninckii (Barrande). Like A. koninckii, the cephalic growth rate in E. kingii was constant and of similar magnitude to the minimum growth rate along the trunk, and growth rates in the trunk were lower during the holaspid ('adult') period than during the meraspid ('juvenile') period. However, body length at the onset of meraspis was smaller, the growth gradient along the trunk during meraspis was shallower, and the terminal number of thoracic tergites was smaller in E. kingii than in A. koninckii. Despite these differences, these two species had similar maximum body lengths, because higher overall growth rates in E. kingii compensated for other differences. The rate of cephalic shape change in E. kingii decreased at the transition from meraspis to holaspis, while the pygidium became more morphologically distinct from the thorax during holaspis. I also provide an emended diagnosis for E. kingii, descriptions of the ontogeny and ventral morphology, and evidence that E. kingii holaspids had an invariant number of tergites. [ABSTRACT FROM AUTHOR]

Trilobite development has been intensively explored during the past decades, but information on ancestral character combinations in the early developmental stages of trilobites remains unknown. Trilobites of the superfamily Olenelloidea are one of the earliest diverging groups. Study of their development coupled with the development of other early diverging trilobite groups can provide information on the ancestral morphology of trilobite early stages. Herein we describe numerous well‐preserved specimens of the olenelloid trilobite Fritzolenellus lapworthi. The earliest stages have circular cephala bearing intergenal spines and lacking genal spines. During subsequent development, morphological changes involve the modification of the cephalic shape from circular to semicircular, expansion of the frontal glabellar lobe, gradual shortening of intergenal spines and origin and prolongation of genal spines. Trunk development of Fritzolenellus suggests that macropleurae and macrospine development are two independent processes and that the origin of the opisthotrunk is linked with the onset of phase 5 of cephalic development. The morphology of the early developmental stages of Fritzolenellus and of some related taxa differs in many aspects from the morphology of equivalent stages of some other members of Olenelloidea. Consequently, two basic morphotypes are recognized during the early development of Olenelloidea: the Fritzolenellus and the Olenellus morphotypes. Comparison with Fallotaspidoidea and Redlichiina indicates that early developmental stages of these taxa share character combinations that are typical for the Fritzolenellus morphotype. Such a comparison suggests that characters defining the Fritzolenellus morphotype are ancestral for Trilobita. The Olenellus morphotype is probably a derived condition within Olenelloidea and might be related to predator deterrence. [ABSTRACT FROM AUTHOR]

Trilobites are one of the most diverse and abundant fossil groups from the early Palaeozoic, and as such are useful for answering important questions about early animal evolution, including developmental processes. Ontogenetic information for a large number of trilobite species has been published, but cases where multiple articulated specimens are known across the full range of developmental stages are rare. The early Cambrian (Series 2, Stage 4) Emu Bay Shale biota from Kangaroo Island (South Australia) is numerically dominated by trilobites, particularly articulated specimens of the ellipsocephaloid Estaingia bilobata, which are present in densities of >600 individuals per square metre on certain bedding planes. Here we describe the essentially complete post‐embryonic ontogenetic series of E. bilobata from the Emu Bay Shale, and investigate patterns of growth relating to articulation and segmentation in this early Cambrian arthropod. Estaingia bilobata exhibits the hypoprotomeric mode of growth, with the epimorphic phase (the cessation of trunk segment generation) reached prior to the onset of the holaspid period. The meraspid pygidium had an extended equilibrium period in which anterior segment release into the thorax was matched by subterminal segment generation. Previously undocumented morphological features of E. bilobata, including the hypostome and bispinose pleural tips in holaspides, are also described. The growth characteristics and morphological features of E. bilobata documented herein strengthen close phylogenetic relationships between the Estaingiidae, Ellipsocephalidae and Xystriduridae. [ABSTRACT FROM AUTHOR]

The new ellipsocephaloid trilobite species Kingaspidoides spinirecurvatus has a spectacular morphology because of a unique set of two long and anteriorly recurved spines on the occipital ring and the axial ring of thoracic segment 8. Together with the long genal spines this whimsical dorsally directed spine arrangement is thought to act as a non-standard protective device against predators. This is illustrated by the body posture during different stages of enrolment, contrasting with the more sophisticated spinosities seen in later trilobites, which are discussed in brief. Kingaspidoides spinirecurvatus from the lower–middle Cambrian boundary interval of the eastern Anti-Atlas in Morocco has been known for about two decades, with specimens handled as precious objects on the fossil market. Similar, but far less spectacular, spine arrangements on the thoracic axial rings are known from other ellipsocephaloid trilobites from the Anti-Atlas of Morocco and the Franconian Forest region of Germany. This suggests that an experimental phase of spine development took place within the Kingaspidoides clade during the early–middle Cambrian boundary interval. [ABSTRACT FROM AUTHOR]

The morphology and ontogeny of the trilobite Changaspis elongata based on 216 specimens collected from the Lazizhai section of the Balang Formation (Stage 4, Series 2 of the Cambrian) in Guizhou Province, South China are described. The relatively continuous ontogenetic series reveals morphological changes, and shows that the species has seventeen thoracic segments in the holaspid period, instead of the sixteen as previously suggested. The development of the pygidial segments shows that their number gradually decreases during ontogeny. A new dataset of well-preserved specimens offers a unique opportunity to investigate developmental traits after segment addition is completed. The ontogenetic size progressions for the lengths of cephalon and trunk show overall compliance with Dyar's rule. As a result of different average growth rates for the lengths of cephalon, trunk and pygidium, the length of the thorax relative to the body shows a gradually increasing trend; however, the cephalon and pygidium follow the opposite trend. Morphometric analysis across fourteen post-embryonic stages reveals growth gradients with increasing values for each thoracic segment from anterior to posterior. The reconstruction of the development traits shows visualization of the changes in relative growth and segmentation for the different body parts. The new dataset and growth gradient of the trunk suggest that the thoracic segment growth dynamics of early Cambrian to Silurian trilobites follow the same general continuous, steady-state growth gradient decreasing from posterior to anterior. [ABSTRACT FROM AUTHOR]

The majority of cornuproetine taxa known from articulated specimens have ten thoracic segments. Here we record rare, well-preserved specimens with nine segments in the Early and Middle Devonian of Morocco, Germany and the Czech Republic. These encompass the genera Lepidoproetus Erben, Sculptoproetus Erben, Macroblepharum Alberti, Koneprusites Přibyl, Paralepidoproetus Alberti, Timsaloproetus Gibb and Chatterton and Hollandiella gen. nov. Large holaspides of these trilobites are often comparatively small and bear dorsal sculpture consisting of terrace lines in a fingerprint-like configuration, small granules, or a mixture of these features. The holotype of Sculptoproetus sculptus (Barrande) and the lectotype of Lepidoproetus lepidus (Barrande), the type species of these genera, are refigured. Additional topotypical specimens of S. sculptus are illustrated; these provide new insights into the morphology of this species, leading to the putative synonymy of Macroblepharum with Sculptoproetus. The following new cornuproetine taxa are described: Hollandiella curvirostris gen. et sp. nov., Hollandiella lebruni gen. et sp. nov., Hollandiella verecunda gen. et sp. nov., Koneprusites oresibios sp. nov., Lepidoproetus arenicolus sp. nov., Sculptoproetus raki sp. nov., Timsaloproetus weddigei sp. nov., Timsaloproetus gibbae sp. nov., Timsaloproetus pulchistriatus pulchistriatus ssp. nov. and Timsaloproetus pulchistriatus repens ssp. nov. Timsaloproetus haasi is recorded from several localities in Morocco. Additionally, Tafilaltaspis rufa sp. nov. is proposed as a new species of the nine-segmented eremiproetine Tafilaltaspis, which was traditionally classified as a cornuproetine genus. [ABSTRACT FROM AUTHOR]

The ontogeny of the pelturine olenid trilobite Leptoplastides salteri (Callaway, 1877) from the Shineton Shales, Shropshire, England, was first described in 1925 by Frank Raw. Since that time, scanning electron microscopy and other new technologies have revealed many more details of structure, of early developmental stages in particular, than were available to Raw. Whereas protaspides are not preserved and the state of preservation is less than perfect for the smallest meraspides, we have established that the latter had an array of delicate, long thoracic and pygidial spines, as well as paired procranidial spines, which disappear by meraspid degree 8. Raw's reconstructions of early meraspides, and his measurements of the early stages in development, are here amended in the light of new information. Dorsal spines in the adult are much more highly developed than have been documented in any other olenid. The hypostome is preserved in place in several specimens. Initially conterminant (attached to the doublure), it becomes natant (free) in late meraspid to early holaspid stages of development, with its anterior contour fitting exactly to that of the glabella. The ecology of the widespread Leptoplastides is best known from very extensive sections in South America, which provide a useful basis for comparison. It was well adapted to a range of environments, both oxygenated and dysoxic, and is usually the dominant taxon in the biofacies in which it is found. [ABSTRACT FROM AUTHOR]

Trilobites are an iconic group of extinct arthropods that lived in Palaeozoic oceans for c. 270 Ma, before their demise at the end of the Permian Period. Despite their considerable diversity (> 22 000 species) and abundance, particularly in Cambrian and Ordovician rocks, as well as intensive study for well over 200 years, trilobite systematics remains in a state of flux. In this contribution, the complex history of trilobite classification over the last century is briefly reviewed, including the seminal scheme published by Henry Swinnerton in 1915. The cryptogenesis problem, which relates to the supposedly obscure phylogenetic links between major post-Cambrian trilobite clades and their Cambrian sister taxa, is also discussed. Previous studies have suggested that the cryptogenesis problem is largely a taxonomic artefact, but the Cambrian origins of some post-Cambrian groups, such as the orders Proetida and Odontopleurida, are still unclear. Future directions for research on trilobite systematics are outlined, from taxonomic studies involving comprehensive documentation and extensive illustration of morphology at the species level, through to broad-scale phylogenetic analyses that initiate or test hypotheses about relationships across the major groups. Other ongoing issues to be addressed include identifying the sister group of Trilobita, and determining whether certain taxa, such as the suborder Agnostina and Cambrian family Burlingiidae, represent trilobites. [ABSTRACT FROM AUTHOR]

Completely enrolled, phosphatized, 3‐D specimens of the eodiscinid trilobite Tsunyidiscus yanjiazhaiensis from Cambrian Stage 3 of South China exhibit much morphological detail and show variation in delicate coaptative structures associated with their trunk segmentation. The relationship between enrolment mechanism and trunk segmentation during ontogeny confirms a unique developmental pattern among the early Cambrian eodiscoid trilobites, revealing how these animals controlled the rate of segment increase and release during post‐embryonic development in their life cycles. [ABSTRACT FROM AUTHOR]

The lower Cambrian (Series 2) White Point Conglomerate (WPC) on Kangaroo Island, South Australia contains exotic clasts representing a diverse array of lithologies, including metamorphics, chert, sandstone, and abundant carbonates, notably archaeocyath-rich bioclastic limestone. Acetic acid digestion of the WPC bioclastic limestone clasts reveals a diverse shelly fauna. This assemblage includes abundant organophosphatic brachiopods such as Cordatia erinae Brock and Claybourn gen. et sp. nov., Curdus pararaensis, Eodicellomus elkaniformiis, Eohadrotreta sp. cf. E. zhenbaensis, Eoobolus sp., Kyrshabaktella davidii, and Schizopholis yorkensis. Additional shelly taxa include the solenopleurid trilobite Trachoparia? sp., the tommotiids Dailyatia odyssei, Dailyatia decobruta Betts sp. nov., Kelanella sp., and Lapworthella fasciculata, spines of the bradoriid arthropod Mongolitubulus squamifer, and several problematica, such as Stoibostrombus crenulatus and a variety of tubular forms. The upper age limit for the WPC is constrained by biostratigraphic data from the overlying Marsden Sandstone and Emu Bay Shale, which are no younger than the Pararaia janeae Trilobite Zone (Cambrian Series 2, Stage 4). The shelly fossil assemblage from the WPC limestone clasts indicates an upper Dailyatia odyssei Zone (= Pararaia tatei to lower P. janeae trilobite zones), equivalent to the Atdabanian-early Botoman of the Siberian scheme. This contrasts with the previously suggested late Botoman age for the limestone clasts, based on the diverse archaeocyath assemblage. The minor age difference between the WPC and its fossiliferous limestone clasts suggests relatively rapid reworking of biohermal buildups during tectonically-active phases of deposition in the Stansbury Basin. [ABSTRACT FROM AUTHOR]

The Olenidae stands out for its abundance and biostratigraphical importance, especially in the Lower Palaeozoic rocks of northwestern Argentina. Their phylogenetic relationships have been traditionally determined stratigraphically and by direct morphological comparison. This study reports the first formal phylogenetic analysis of olenids. Eighty‐six characters (24 quantitative and 62 qualitative) were coded for 65 taxa (58 olenids). Quantitative characters were treated both as discrete and as continuous variables. To explore the best way of character coding for this group, continuous characters were coded as: median, log‐median, normalized and rescaled. Maximum parsimony and implied weighting were used as optimality criteria. A phylogenetic hypothesis more consistent with traditional taxonomy was reconstructed with both quantitative and qualitative partitions. All the trees obtained with quantitative characters coded as continuous and rescaled are better resolved, and those topologies were more similar among them. This treatment also reflects more effectively the behaviour of the original variables. Olenidae is not a monophyletic clade: Andrarina costata and Aphelaspis australis are included within the ingroup, as sister clade of Olenus gibbosus. Also, the results suggest that members of the Hypermecaspidinae constitute a new family within the Order Olenida. The traditional taxonomic scheme at subfamily level is partially supported. Triarthrinae and 'pelturinds' are recovered as monophyletic clades, but Oleninae is polyphyletic. This study proves, through a formal cladistic analysis, that characters disregarded by traditional taxonomy can be uncovered. Finally, this is the first step towards achieving a classification of the Olenidae taking into account the evolutionary process involved in its diversification history. [ABSTRACT FROM AUTHOR]

Two hundred and eighty specimens of paradoxidids from two localities in the Holy Cross Mountains (Poland) have been reanalysed using morphometric techniques. Revision of the dataset provided evidence for the presence of two endemic taxa: Acadoparadoxides kozlowskii (Orłowski, 1959) and Acadoparadoxides samsonowiczi (Orłowski, 1959), both belonging to the earliest group of Acadoparadoxides , initially considered to be present only in Gondwanan successions. Thus, this is the first description of the members of this group outside West Gondwana. The taxonomic revision, and the presence of representatives of the earliest acadoparadoxidines coupled with the absence of taxa typical of Scandinavia that were previously described from this locality have resulted in the modification of the biostratigraphic scheme hitherto used in the Holy Cross Mountains. The newly established A. samsonowiczi – A. kozlowskii Assemblage Zone is correlated with the Morocconus notabilis Zone of Morocco, and thus represents the Cambrian Series 2 and 3 boundary interval. Links with West Gondwana challenge the existing palaeogeographic interpretations for the southern part of the Holy Cross Mountains and point to an urgent need to revise the position of the Małopolska Block during the middle Cambrian. [ABSTRACT FROM AUTHOR]

Euarthropods have a tough exoskeleton that provides crucial protection from predation and parasitism. However, this is restrictive to growth and must be periodically moulted. The moulting sequence is well-known from extant arthropods, consisting of: (i) the long inter-moult stage, in which no changes occur to the hardened exoskeleton; (ii) the pre-moult stage where the old exoskeleton is detached and the new one secreted; (iii) exuviation, when the old exoskeleton is moulted; and (iv) the post-moult stage during which the new exoskeleton starts as soft, thin, and partially compressed and gradually hardens to the robust exoskeleton of the inter-moult stage. Trilobite fossils typically consist of inter-moult carcasses or moulted exuviae, but specimens preserving the post-moult stage are rare. Here we describe nine specimens assigned to Symphysurus ebbestadi representing the first group of contemporaneous fossils collected that preserve all key stages of the moulting process in one taxon, including the post-moult stage. They were collected from a single lens in the Tremadocian part of the Fezouata Shale Formation, Morocco. Based on cephalic displacement and comparison to other trilobite moults, one specimen appears to represent a moulted exoskeleton. Four specimens are typical inter-moult carcasses. Four others are wrinkled and flattened, with thin exoskeletons compared to inter-moult specimens, and are considered post-moult individuals. These S. ebbestadi specimens illuminate the preservation and morphology of the post-moulting stage, characterised by strong anterior-posterior exoskeleton wrinkling, as well as overall body flattening and reduced visibility of thoracic articulations. Being found in the same lens, these specimens likely represent the first preserved in-the-act mass moulting event. The displayed sequence of moulting suggests the moulting process in trilobites was comparable to modern arthropods, and conserved within euarthropod evolutionary history. [ABSTRACT FROM AUTHOR]