Late Ordovician brachiopods from eastern North Greenland: equatorial offshore migration of the Red River fauna

Abstract: Late Ordovician rhynchonelliformean brachiopods, typical of the North American Red River fauna, are found sporadically in the Børglum River Formation of the Centrum Sø area, Kronprins Christian Land, eastern North Greenland. The geographical distribution of this characteristic brachiopod fauna is thus extended to the easternmost extremity of the Laurentian craton. The assemblage compares specifically with the Hiscobeccus brachiopod fauna, based on key taxa such as notably Hiscobeccus gigas (Wang, 1949), and indicates a late Katian age for this part of the succession. For the first time, this typically inland, shallow‐water fauna is found associated with genera like Bimuria, suggesting a transitional marginal facies with outer shelf benthos. The current study describes a Hiscobeccus fauna that lived at the seaward edge of its preferred habitat. Furthermore, an unpublished Hiscobeccus fauna, from the Børglum River Formation of Peary Land, central North Greenland, as well as several occurrences from the Kap Jackson and Cape Calhoun formations in various parts of Washington Land, western North Greenland, are described here. These show a distinct shift from older strata containing H. capax (Conrad, 1842) to younger strata exclusively yielding specimens of H. gigas. As H. gigas occurs in the upper part of the Cape Calhoun Formation in Washington Land, it indicates that the upper boundary of the Cape Calhoun Formation is considerably younger than previous estimates, reaching into the uppermost Katian (middle Cautleyan–Rawtheyan). The Cape Calhoun Formation correlates with the upper member of the Børglum River Formation and further demonstrates that the Hiscobeccus fauna was widespread in Laurentian marginal settings of North Greenland. Even though the Hiscobeccus fauna was pan‐continental during the late Katian (Richmondian), it possesses a strong provincial signal during the later Ordovician. The new occurrences indicate that this fauna extended to the north‐eastern margin of the Laurentian Craton. It lived in close association with cosmopolitan faunal elements that may have been the earliest sign of the succeeding invasion of migrants from Baltica that arrived later during the Hirnantian. The offshore migration of this atypical Hiscobeccus fauna likely demonstrates the path of warm‐water currents as the Centrum Sø locality was located at the equator during the Late Ordovician.     

Katian (Upper Ordovician) conodonts from Wales

Middle and upper Katian conodonts were previously known in the British Isles from relatively small collections obtained from a few localities. The present study is mainly based on 17 samples containing more than 17 000 conodont elements from an approximately 14‐m‐thick succession of the Sholeshook Limestone Formation in a road cut near Whitland, South Wales, that yielded a diverse fauna of more than 40 taxa. It is dominated by representatives of Amorphognathus, Aphelognathus/Plectodina and Eocarniodus along with several coniform taxa. Representatives of Decoriconus, Istorinus and Sagittodontina are reported from the Ordovician of UK for the first time. The fauna is a typical representative of the British Province of the Atlantic Realm and includes a mixture of taxa of North American, Baltoscandic and Mediterranean affinities along with pandemic species. Based on the presence of many elements of Amorphognathus ordovicicus and some morphologically advanced specimens of Amorphognathus superbus, the Sholeshook Limestone Formation is referred to the lower A. ordovicicus Zone. Most of the unit is also coeval with Zone 2 of the Cautleyan Stage in the British regional stage classification, and stage slice Ka3 of the middle Katian Stage in the global stratigraphical classification, an age assignment consistent with data from trilobites, graptolites and chitinozoans. The unusually large collection of M elements of Amorphognathus provides insight into the complex morphological variation in this element of some Katian species of this genus. The Sholeshook conodont fauna is similar to those of the Crûg and Birdshill limestones, but differs in several respects from the slightly older ones from the Caradocian type area in the Welsh Borderland. Although having some species in common, the Sholeshook conodont fauna clearly differs from coeval Baltoscandic faunas and is even more different in composition compared with equivalent North American Midcontinent faunas.     

Conodont biostratigraphy and carbon isotope chemostratigraphy of the Middle to Upper Ordovician on the western Yangtze Platform,South China

Middle and Late Ordovician strata in the nearshore area of the western Yangtze Platform are investigated. In total, 241 δ¹³C_carb samples and 108 conodont samples from the Songliang and Tuanjie sections in the northeastern Yunnan Province were collected for chemostratigraphic and biostratigraphic studies. Altogether, 21 species belonging to 16 conodont genera were recovered from the two sections, among which Ansella jemtlandica, Drepanoistodus basiovalis, Hamarodus brevirameus, and Microzarkodina hagetiana are important age indicators within the Dapingian to early Katian interval. Four carbon isotope shifts are recognized in the studied sections: a negative shift in the early Darriwilian, the middle Darriwilian carbon isotope excursion (MDICE), the early Katian Guttenberg carbon isotope excursion (GICE), and a further positive carbon isotope shift later in the early Katian which is a new record of this event in South China. Integrated carbon chemostratigraphy and conodont biostratigraphy indicate early Darriwilian and early Katian ages for the base and top, respectively, of the Huadan Formation at the Songliang section, and a Darriwilian to early Katian age for the Shihtzupu Formation at the Tuanjie section.     

Discovery of Anticostia uniformis from the Xiazhen Formation at Zhuzhai,South China and its stratigraphic implication

The Xiazhen Formation is an Upper Ordovician lithostratigraphic unit in the Jiangshan-Changshan-Yushan (JCY) area, which contains series of Late Ordovician reef successions. The reef successions of the Xiazhen Formation at Zhuzhai are critical for evaluation of the Late Ordovician marine diversity and palaeoecology. However, their age has long been uncertain and generally is regarded as of upper Katian, based on the occurrences of shelly fossils and correlation with the stratigraphic equivalent Changwu Formation. The newly discovered graptolite species Anticostia uniformis, in the Xiazhen Formation, together with the combined evidence of brachiopods and sedimentology, indicates an age range for the graptolite locality from the Dicellograptus complanatus Biozone to the Diceratograptus mirus Subzone of late Katian, but the graptolites do not rule out the possibility that it is Hirnantian.     

Upper Katian (Upper Ordovician) trans-Atlantic δ13C chemostratigraphy: the geochronological equivalence of the ELKHORN and PAROVEJA excursions and its implications

Since 2010 when the North American ELKHORN and Baltoscandic PAROVEJA isotope excursions were first described and named, their mutual age relations have remained uncertain, if not controversial. This was at least partly due to the incompleteness of the ELKHORN excursion in its reference section in western Ohio. The unexpected discovery of an apparently complete ELKHORN excursion in a drill core from St Marys in western Ohio has led to the conclusion that in terms of stratigraphical position and δ¹³C curve correspondence, the ELKHORN and PAROVEJA excursions are so similar that they apparently represent the same isotopic curve perturbation. The ELKHORN/PAROVEJA excursion occurs in the D. pacificus Graptolite Zone and uppermost A. ordovicicus Conodont Zone in the uppermost Katian Stage (Stage Slice Ka4 of Bergström et al. Lethaia 42, 97–197, 2009). Because the designation PAROVEJA was published two months before that of ELKHORN, it has priority as excursion designation. This excursion is particularly well represented in the carbonate successions in the Great Basin of western United States. Chemostratigraphy and biostratigraphy in that region show that the Richmondian transgression was contemporaneous with the beginning of the middle Katian WHITEWATER/MOE excursion. The onset of the Richmondian transgression has long been controversial but now available evidence suggests that it is of essentially the same age across large regions of the southern, western and central United States.     

Mosasaur predation on a nautiloid from the Maastrichtian Pierre Shale,Central Colorado,Western Interior Basin,United States

Mosasaurs were common predators on the ammonites that inhabited the upper water column of the Late Cretaceous Western Interior epicontinental seaway of North America. Mosasaurs developed predictable behaviour patterns for feeding on ammonite prey. There are no previous reports of mosasaur predation on the much less common Cretaceous nautiloids, possibly because of the prey's predominantly deep, epibenthic habitat, as deduced from modern Nautilus life habits. A single specimen of the highly inflated nautiloid, Eutrephoceras dekayi (Conrad), prey to a small adult mosasaur, likely Platycarpus, Prognathodon or Mosasaurus, is reported herein from the Pierre Shale of Colorado in the Early Maastrichtian biozone of Baculitesgrandis transitional to the biozone of B. clinolobatus. The nautiloid was killed in the same manner as described previously for discoid ammonites (Placenticeras, Sphenodiscus) from coeval strata in the USA and Canada.     

Caradoc strophomenoid and plectambonitoid brachiopods from Wales and the Welsh Borderland

Abstract: Ordovician strophomenoidean and plectambonitoidean brachiopods are reviewed and partially redescribed from the Caradoc (Sandbian to Early Katian) age rocks of Wales and the Welsh Borderland of England, then forming part of the Avalonia Terrane. There are 51 nominal species available, of which 12 are synonymised here and many transferred to different genera. Strophomenoids occurred sporadically, often rarely but occasionally abundantly, particularly in mid‐shelf benthic assemblages, and 21 named species and subspecies are noted, including the new Kiaeromena (Kiaeromena) harperi, as well as six in open nomenclature. The common genus Kjaerina, with six species known from the area, is revised in detail. In contrast to the strophomenoids, plectambonitoids were more common in shallow and mid‐shelf assemblages, including Eoplectodonta (Eoplectodonta) abigailae sp. nov., with Sowerbyella in particular dominating many bedding planes, and 17 named species are reviewed, as well as three in open nomenclature. The distinctive Gunningblandella, hitherto known only from Australia and Kazakhstan, is recorded for the first time from Europe. The Avalonian strophomenides are compared with those from neighbouring terranes, particularly Laurentia and Baltica, during a time when the seas between those terranes were narrowing, and the faunas from the three terranes are found to be largely similar at the generic level, although there are few species in common. In contrast, there are few genera in common between those three terranes and the Mediterranean Province of Northwestern Gondwana.     

A new ichthyodectiform (Actinopterygii,Teleostei) from the Lower Cretaceous Marizal Formation,north‐east Brazil

Abstract: A new genus and species, Ogunichthys triangularis, from the Early Cretaceous Marizal Formation in the Tucano Basin, Bahia State, Brazil, can be assigned to the teleost order Ichthyodectiformes and to the suborder Ichthyodectoidei by the presence of ten synapomorphies of these clades. Ogunichthys gen. nov. cannot be placed into any of the currently recognized families within Ichthyodectoidei and differs from all other ichthyodectoid genera in the ratio between the depth of the dentary symphysis and the depth of the coronoid process, the relationships between the length of the coronoid process and the alveolar border of the dentary, and a supraoccipital crest with posterior border fringed. The new taxon occurs in Early Cretaceous (Aptian) strata of the Marizal Formation, which are related to periods during epicontinental seas extended from the Caribbean Tethys into the South American continent. Rather than representing a single continuous widespread seaway, these epicontinental seas reflect local tectonic events, sometimes including locally distinctive faunas.     

The Middle–Late Ordovician brachiopod Plectorthis from North America and its paleobiogeographic significance

Plectorthis is a commonly reported Late Ordovician brachiopod genus from Laurentia and several other tectonic plates. Examination of shells from the Cincinnati Arch type area shows that species previously assigned to the genus have a considerable range of variability, such as ribbing style, shell outline, and musculature. The type species, Orthis plicatella Hall, 1847 is characterized by generally simple costae, with double-rowed aditicules on each rib, and a cordate ventral muscle field. Some species previously attributed to Plectorthis have a subtriangular ventral muscle field and hence a closer affinity to the plaesiomyid genus Austinella. Orthis fissicosta is here demoted to a subspecies of Plectorthis plicatella based on its similar ribbing style to some specimens of the type species. In Laurentia, Plectorthis first appeared in the Darriwilian–Sandbian boundary interval, achieved a relatively high species diversity and wide paleogeographic distribution in pericratonic seas during the early Katian; the subequatorial species attained a notably larger shell than those in the higher paleotropics. The genus did not invade the intracratonic basins of Laurentia and became extinct by the late Katian.     

Upper Ordovician bryozoans from the Xiazhen Formation of Yushan,northeastern Jiangxi,East China

The bryozoan fauna from the Xiazhen Formation (Katian, Upper Ordovician) of northeast Jiangxi Province, southeast China is reported here. Seventeen species of bryozoans belonging to fifteen genera and four orders are identified: Homotrypa yushanensis, Homotrypa sp., Prasopora yushanensis, Trematopora sp., Monotrypella sp., Rhombotrypa sp., Orbignyella sp., Constellaria jiangxiensis, Constellaria sp., Stictopora nicholsoni, Trigonodictya parvula, Ptilodictya ensiformis, Stictoporella sp., Pseudopachydictya sp., Nematopora sp., Arthrostylidae sp. indet., and Chasmatoporidae sp. indet. Four of these genera have been reported previously but nine genera (Trematopora, Monotrypella, Rhombotrypa, Orbignyella, Trigonodictya, Ptilodictya, Stictoporella, Pseudopachydictya, and Nematopora), one rhabdomesine and one fenestrate are found for the first time in the Late Ordovician strata of South China. Our palaeogeographical analysis suggests that the bryozoan association is typical for the Katian, which is mostly widespread in Laurentia, Siberia, Baltica and Mediterranean, and displays palaeobiogeographical relationships to the Laurentia–Siberia Province.     

Genetic and morphological evidence for cryptic diversity in the <Emphasis Type="Italic">Careproctus rastrinus</Emphasis> species complex (Liparidae) of the North Pacific

The molecular phylogeny of the Careproctus rastrinus species complex is presented on the basis of sequence variations in the 16S rRNA and cytochrome b genes (1,447 base pairs) of mitochondrial DNA using specimens collected from across the North Pacific and its marginal seas, including the Sea of Japan, the Pacific coast of Japan, the Sea of Okhotsk, the Bering Sea, the Gulf of Alaska, and the Arctic Ocean. Gene flow and migration between geographic regions is apparently strictly restricted in the C. rastrinus species complex, as this phylogeny revealed nine genetically diverged groups in the C. rastrinus species complex, most of which corresponded well with major geographic regions. Most of the groups were different in terms of morphological characters. Five nominal species have been considered to be members of the complex and have been variously recognized as being valid or as synonyms of C. rastrinus, but the present genetic and morphological differences suggested that the C. rastrinus species complex represents far more divergence. Such cryptic diversity of the C. rastrinus species complex defined by geographic regions may be related to their low dispersal ability, because they bear large demersal eggs. The genetic divergence suggested that colder climates from the late Pliocene and the isolation of marginal seas during the Pleistocene may have driven the divergence of the C. rastrinus species complex.     

Middle–Late Ordovician chitinozoans from Songliang of Qiaojia,western South China,and their biostratigraphic implications

Ordovician chitinozoans are as yet inadequately documented from the western Yangtze Platform, South China. Here we present a systematic study on chitinozoans from a Middle–Upper Ordovician succession at Songliang of Qiaojia, northestern Yunnan, southwestern China. Altogether 34 species of 12 genera are identified from the upper Hungshihyen and the Huadan formations. The top of the Hungshihyen Formation yields typical Early and Middle Ordovician forms, for instance, Lagenochitina obeligis and Belonechitina chenjiawuensis, with the latter only known from the lower Darriwilian in South China, thus suggesting an early Darriwilian age for this interval. The chitinozoans recovered from the Huadan Formation include Lagenochitina prussica and Spinachitina fossensis, both are mainly confined in Katian and sometimes ranging into younger strata. This indicates an age younger than the middle to late Darriwilian previously determined for the Huadan Formation. However, conflicts exist in age assignment of the Huadan Formation based on chitinozoan and other evidence, and further work is required.     

Cincinnetina,a new Late Ordovician dalmanellid brachiopod from the Cincinnati type area,USA: implications for the evolution and palaeogeography of the epicontinental fauna of Laurentia

Abstract: The most common forms of Late Ordovician dalmanellid brachiopods from the Cincinnatian type area, previously treated as either Dalmanella or Onniella, are assigned to Cincinnetina gen. nov. The new genus differs from Dalmanella and Onniella in having a consistently developed primary medial costa in the dorsal valve, a larger cardinal process that tends to develop a trilobed myophore, strongly differentiated fine and coarse punctae, and sparse aditicules. Cincinnetina can be distinguished from the closely related Paucicrura and Diceromyonia in its smaller trilobed cardinal process (when developed) that does not have a dominant medial lobe and does not extend into the delthyrial cavity of the ventral valve. Globally, Dalmanella and Onniella occur most commonly in deposits of relatively deep‐ or cool‐water palaeoecological settings, whereas in North America, Cincinnetina, Paucicrura and Diceromyonia are found mainly in carbonate‐rich deposits in warm‐water depositional environments, with Cincinnetina and Paucicrura most common in pericratonic settings and Diceromyonia in palaeoequatorial inland seas.     

A reassessment of the systematics and a proposal for the phylogeny of some cosmopolitan Lineus species (Nemertea)

Morphological, behavioural and biochemical characters of 9 cosmopolitan, benthic species of the nemertean genus Lineus are used both to clarify their taxonomy and to discuss their phylogenetic relationships. Data analysis shows that all fissiparous species of Lineus collected in the world seas up to date can be placed in a single species, L. sanguineus, with three recognisable subspecies, L. s. sanguineus, L. s. nigricans and L. s. pseudolacteus. These taxa were originally described as separate species mainly according to their geographic origin under the names L. sanguineus (shores of European seas), L. socialis (Atlantic North American shores), L. vegetus (Pacific North American shores), L. pseudolacteus (shores of the English Channel) and L. nigricans (Mediterranean coasts of Italy and France). Examination of the character value matrix suggests a phylogenetic tree developing from two ancestral branches. The first branch includes the two related species L. ruber and L. viridis, the second branch the three species L. longissimus, L. lacteus and L. sanguineus. This approach to the taxonomic and phylogenetic relationships of Lineus nemertean species is in agreement with the intra- and interspecies histocompatibility data in nemerteans: (i) grafts transplanted from donors to recipients of the same species succeed and grafts transplanted from donors to recipients of different species fail; (ii) grafts are rejected more rapidly when the donor and recipient species diverged earlier in evolution and, a contrario, grafts are accepted better when the donor and recipient species have been more recently isolated.     

Carboniferous biogeography of the bryozoan Archimedes in North America

The fenestrate bryozoan genus Archimedes apparently originated and then diversified in Early Carboniferous seas of eastern North America. Its highest species richness and greatest abundance are reached in uppermost Lower to lowest Upper Carboniferous (Chesterian) rocks of eastern North America. Following the unconformity that marks the mid‐Carboniferous eustatic event, however, it is known in North America only in Upper Carboniferous basinal deposits of the Oquirrh Mountains and in nearby regions of Utah and Nevada. It is hypothesized that Archimedes’ life‐history patterns caused the biogeographical change: dependence on fragmentation as a means of recruitment did not allow Archimedes to keep up with its rapidly migrating preferred back‐barrier environment during the numerous eustatic transgressive and regressive cycles that caused the shoreline to sweep across North America during the Late Carboniferous. The persistence of Archimedes in the Oquirrh Basin and vicinity is inferred to have resulted from its preferred back‐barrier environment migrating only locally along the relatively steep basinal slope present there during the Late Carboniferous eustatic cycles.     

STROPHOMENIDE BRACHIOPODS FROM THE CHANGWU FORMATION (LATE KATIAN, LATE ORDOVICIAN) OF CHUN'AN, WESTERN ZHEJIANG, SOUTH-EAST CHINA

Abstract:  Late Ordovician strophomenide brachiopods (superfamilies Strophomenoidea and Plectambonitoidea) from the upper Changwu Formation (mid Ashgill, late Katian) of Jianglütang, Chun'an County, western Zhejiang Province, consist of ten genera and 12 species. Five new species of three new genera are recognized: Chunanomena triporcata , Chunanomena sembellina , Cheramomena subsolana , Lateriseptomena modesta , and Lateriseptomena rugosa . The strophomenide brachiopods from the upper Katian strata described in this study and those from the border region of Zhejiang and Jiangxi provinces reported in previous work contain 16 strophomenoid and 12 plectambonitoid genera, and most of the strophomenoids are endemic to South China. Numerical analysis of well-documented late Katian strophomenide brachiopod faunas indicates a strong provincialism, characterized by the highly distinct North American province (Laurentia), the South China-Kazakhstan province, and the Avalonia-Baltica province (Wales, Belgium and Sweden). Surprisingly, the Girvan district of Scotland, which was a peri-Laurentian terrane during the Ordovician, contains a late Katian brachiopod fauna that is more closely related to the contemporaneous brachiopods of Avalonia-Baltica than to those of North America.     

Stratigraphy of the Lungmachi black shales from the Huaying drill core in southwestern China and its palaeogeographic implications

Core samples from the Huaying well, Sichuan Province, have been analyzed for both graptolite biostratigraphy and carbon isotope chemostratigraphy. The result shows a stratigraphic sequence of Lituigraptus convolutus, Stimulograptus sedgwickii and Spirograptus guerichi biozones, and records a strong positive shift of δ¹³C_org at the base of the Stimulograptus sedgwickii Biozone, which coincides with those from Cornwallis Island (Arctic Canada), Arisaig (Nova Scotia, Canada), and the Barrandian area (Bohemia). Two outcrops were also studied in the Huaying region. The Yanwanggou section shows a continuous, but complicated, stratigraphical sequence, from the upper Katian to the lower–middle Rhuddanian, then a sedimentary hiatus of ∼3.26 Ma, and finally a continuous sequence from the upper Aeronian to the lower Telychian, whereas the Liziya section and the Huaying well exhibit significant hiatus, which lasted from Hirnantian to middle Rhuddanian or early Aeronian times. Therefore, the eastern boundary of the Central Sichuan Oldland during the Ordovician and Silurian transition can be precisely constrained to a narrow area between the Huaying well-Liziya section and the Yanwanggou section, which indicates that the Central Sichuan Oldland was much larger than previously considered. The present data also indicate the existing of a so far unidentified stage of the Kwangsian Orogeny, which resulted in the uplift of some portions of the Yangtze Platform region and the consequent delay of the transgression in those places. The multi-stage hiatus, spanning at least from the upper Katian to middle Aeronian in the study area, resulted from a combined effect of the Hirnantian glacio-eustatic sea-level fall and the regional tectonic activity of the Kwangsian Orogeny.     

Oldest known Dicoelosia and Epitomyonia,deep water brachiopods from the Beiguoshan Formation (Middle Katian,Upper Ordovician), Shaanxi,north China

Abstract: A diverse brachiopod fauna from a relatively deep water carbonate facies of the Upper Ordovician Beiguoshan Formation (uppermost Caradoc – lower Ashgill, middle Katian) is characterized by small shells and contains the oldest known Dicoelosia and Epitomyonia, two diagnostic taxa of deep water brachiopod palaeocommunities during the Late Ordovician and Silurian. Three new species are recognized: Dicoelosia cordiformis sp. nov., Dicoelosia perbrevis sp. nov. and Epitomyonia fui sp. nov. These pioneer forms of the family Dicoelosiidae show a relatively high degree of morphological plasticity. The shells of Dicoelosia from the Beiguoshan Formation range from the typical slender‐lobed form with a concavoconvex profile to the strongly equibiconvex, fat‐lobed morphotype that was not known previously until the late Silurian. The Beiguoshan dicoelosiids point to an important attribute of the deep water brachiopods: small generalists with high morphological plasticity, which make them ideal candidates as progenitors for the evolution of shallow water brachiopod faunas in shelf and platform depositional environments.     

Scutelluid trilobites from the Silurian of New South Wales

Abstract: Twenty‐six species (nineteen of them formally named) of scutelluid trilobites belonging to eight genera are described from limestones of the lower Wenlock to Ludlow Mirrabooka Formation and its stratigraphical equivalents in the Orange district of New South Wales. Two genera, Borenoria and Mirrabooka, are new. Japonoscutellum, previously employed mainly for several species from Japan, is a globally widely distributed genus in rocks of middle Llandovery to Ludlow age and includes a number of species previously assigned to Kosovopeltis. Japonoscutellum is remarkably diverse in the faunas from Orange where it is represented by eleven species, one of them known also from Kazakhstan. Flexiscutellum and Tosacephalus are regarded as junior synonyms of Decoroscutellum and Kosovopeltis, respectively. The homology of various scutelluid cephalic structures is discussed, in particular the cephalic borders and border furrows that have routinely been misidentified; the term ‘bolus’ is introduced for the variably developed swelling that is enclosed by S1 on the glabella of most scutelluids. New species are Australoscutellum iota, A. microps, A. pulex, A. trica, Borenoria cirrita, Decoroscutellum planes, Eoscutellum annosum, Illaenoscutellum tryo, Japonoscutellum borenorense, J. crassum, J. diascetum, J. gephyricum, J. guttulatum, J. serangicum, J. strabo, Kosovopeltis trepida, K. xynon and Mirrabooka harryi. Some of these trilobites show strong affinity with species from the Kurosegawa Terrane of south‐western Japan and the Chu‐Ili Terrane of Kazakhstan, as previously reported also for the effaced Illaenina from the same limestones. Australoscutellum and Mirrabooka are presently known only from eastern Australia. The new genus Securifrons is proposed for several species previously assigned to Kosovopeltis or Eokosovopeltis from the Sandbian or Katian to Llandovery of North Greenland, Canada, Scotland, South China and New South Wales.