A NEW SPECIES OF CYRTOSPIRIFER (BRACHIOPODA) FROM THE MIDDLE DEVONIAN OF THE WESTERN SAHARA (NORTHWESTERN AFRICA)

A new species of Cyrtospirifer is described from the Middle to Upper Givetian of the Western Sahara (Northwest Africa). Cyrtospirifer tindoufensis new species differs in its smaller number and coarser medial and flank plications and equibiconvex shell profile from the other Givetian species of Cyrtospirifer that all occur in Europe and to which the new species probably gives rise. The new implications of the proposed phylogeny of the earliest cyrtospiriferids and their origin from the Western Sahara are discussed. The palaeogeographic distribution of the cyrtospiriferids during the Givetian and Frasnian is shown and its migration ways are described considering the global transgression and regression cycles.

New data on internal morphology of exceptionally preserved Nannirhynchia pygmaea (Morris, 1847) from the Lusitanian Basin (Brachiopoda, Early Jurassic, Portugal)

Pyritized internal moulds of articulated shells of the Early Jurassic brachiopod taxon <i>Nannirhynchia pygmaea</i> were found in beds closely below the early Toarcian oceanic anoxic event in the <i>Polymorphum</i> Zone in Portugal. The material allows a detailed study of the outline of the muscle fields, the length and direction of the crura, and the orientation of the cardinalia, which are hitherto undescribed. Three-dimensional reconstructions of articulated shells of <i>N. pygmaea</i> occurring in a single horizon were produced to show the orientation and length of arcuiform crura. The preservation of internal moulds together with the three-dimensional reconstruction of the internal shell morphology allow a more precise description of the internal morphology of this taxon than it is possible with articulated shells and serial sections. <br><br> doi:<a href="http://dx.doi.org/10.1002/mmng.201200005" target="_blank">10.1002/mmng.201200005</a

A Jurassic (Bathonian-Callovian) Daghanirhynchia Brachiopod Fauna from Jordan

A Jurassic (Bathonian-Callovian) brachiopod fauna from Jordan consists of seven rhynchonellid species all belonging to the genus Daghanirhynchia of which two are new: Daghanirhynchia rawyaensis and D. jordanica. Emended diagnoses are given for Daghanirhynchia daghaniensis and D. macfadyeni. Additional taxa described include Daghanirhynchia angulocostata, D. susanae and D. triangulata. Threedimensional reconstructions illustrate the internal morphology of the articulated shells for the first time in this genus. The material studied herein was collected from Wadi Zarqa in northwestern Jordan, almost due north of the Dead Sea, and to the east of the Rift Valley. Most species seem to be geographically restricted within the Jurassic Ethiopian Province, however specimens from Somalia and Ethiopia are larger in size than in other parts of the Province and shell size increases in stratigraphically younger specimens. The occurrence of Daghanirhynchia in India is the only appearance of the genus outside the Ethiopian Province

Taphonomy and palaeoecology of the green Devonian gypidulid brachiopods from the Aferdou El Mrakib, eastern Anti-Atlas, Morocco

On Aferdou El Mrakib, a large reef mound in the Maïder region (Anti-Atlas, Morocco), thick-shelled gypidulids of two genera are locally very abundant. Like Stringocephalus in the shallow water limestone formations in Germany, these Moroccan brachiopods of the genera Devonogypa and Ivdelinia often display greenish shells. By analysing these shells by EDX, it turned out that the colour was possibly caused by impurities of Fe2+-ions. The concentration varies, indicating that the colour is less dependent on the concentration than on shell thickness, because only the thickest parts of the shells appear green and thin-shelled forms never display the green colour. There is also some indication that the Fe content increases towards deeper shell layers (further away from the surface). In addition, we examined the quality and spatial distribution of sublethal injuries in over 200 specimens of Devonogypa and Ivdelinia. Shape, spatial distribution on the shells, and abundance of the sublethal injuries support the hypotheses that (1) the injuries had several causes, (2) some of these were inflicted by predators, probably cephalopods, and (3) many fractures and deformations might have been caused by the brachiopod shells hitting each other in dense populations in agitated water. The existence of dense clusters, built by the association of members of both genera or of only one taxon, is corroborated by the patchy occurrence of these brachiopod

Mapa Geológico de España E 1:200000, hoja 69, Pozoblanco

Peer reviewe

Data from: A new Givetian Athyridid species from NW Africa discovered by 3D reconstruction of shell morphology of internal molds

A new method of analyzing the internal shell morphology (including the complete brachidium of internal molds) of articulated brachiopod shells through the use of serial sections and digital three-dimensional reconstruction is presented. The method introduced is essential for the study of internal shell structures such as brachidia, or cardinalia, if computed tomography (CT) is not suitable or if a CT scanner is not available. A new species of Athyris from Givetian beds of Northwest Africa was selected to exemplify this method. To compare this species with figured serial sections in the literature, two-dimensional drawings of grinding surfaces are provided. Athyris africana n. sp. is only preserved as internal molds of articulated specimens. The new species is included in the evolutionary lineage of the group around Athyris concentrica. The faunal assemblage of A. africana n. sp. shows affinities to Western European and North American brachiopod faunas

Supplemental Data 2

3-D reconstruction of Athyris africana n. sp. created in SPIERSedit based on Supplemental Data 1. The reconstruction provided is weakly smoothed

Supplemental Data 3

Measurements of material studied. Abbreviations: *=holotype; ●=figured; ∆=drawn; +=sectioned; ◊=SEM

Phylogeny, taxonomy and palaeobiogeography of delthyridoid Spiriferids (Brachiopoda, Silurian to Devonian)

Delthyridoid spiriferids are characterized by a global abundance and fast evolution during Silurian and Devonian, and, therefore, are used as important biostratigraphical and palaeobiogeographical tools. In this work, delthyridoid brachiopod faunas from different regions of today’s world, resp., of different palaeobiogeographical units, are compared side-by-side to investigate their phylogenetic relationships and to improve, in a second step, the palaeobiogeography from Late Silurian to Early Eifelian time. A new systematics of Delthyridoidae is established which is more complicated than hitherto assumed. The results of this study are mainly based on direct comparison of articulated and isolated brachiopod shells, external and internal moulds, as well as latex casts and serial sections. The computer supported cladistic analyses have turned out not to be useful due to different kinds of preservation resulting in an incomplete matrix which is insufficient for reliable cladograms. A further problem in terms of cladistical analyses are various convergences during the evolution of spiriferids. Many characters evolved independently from each other at different times in each lineage so that autapomorphies are hardly or not at all recognizable. As a result, families and genera are only definable by a combination of characters rather than by a single or a few autapomorphies. As a new method, 3D reconstruction from serial sections is introduced which made it possible for the first time to compare directly mouldic and shelly material. Preliminary results are presented herein. Statistical analyses of measurements taken from new taxa are made but regarded as a descriptive argument rather than a deciding factor for taxonmy due to incomplete preservation and/or tectonic deformation. Brachiopods, especially type material, from collections of different institutions and museums are studied as well as personal material, whenever possible collected from topotype outcrops. Emended diagnoses, if necessary, from family to species level are given. During this work several new taxa have been erected: 7 new families: Australospiriferidae, Murchisonispiriferidae, Orientospiriferidae, Otospiriferidae, Patriaspiriferidae, Rostrospiriferidae, and Trigonospiriferidae; 6 new genera, 1 of these in open nomenclature: Cyclopterospirifer, Hallispirifer, Parlinispirifer, Murchisonispirifer, Shujiapingensispirifer, and gen. nov. B; and 3 new species: Patriaspirifer merriami, Patriaspirifer johnsoni, and Murchisonispirifer feldmani; 1 taxon is defined as nomen novum: Orientospirifer nakaolingensis wani. In the framework of this project, 2 families: Filispiriferidae and Multispiriferidae; 1 subfamily: Multiplicatispiriferinae, 6 genera, 1 of them in open nomenclature: Frequentispirifer, Leonispirifer, Multiplicatispirifer, Ovetensispirifer, Turcispirifer, and Gen. A; and 9 new species, 3 of them in open nomenclature: Filispirifer hamadae, Leonispirifer leonensis, Multiplicatispirifer foumzguidensis, Oventensispirifer novascotianus, Quiringites arensentiae, Turcispirifer turciae, Multiplicatispirifer cf. foumzguidensis, Quiringites cf. arensentiae, and ?Turcispirifer sp. A which have already been established are also described in this work. The brachiopod faunas studied consist of externally very similar spiriferids which have been identified as same genera, species, or even subspecies in earlier times. These forms are considered as 6 distinct morphotypes Howellella-, Arduspirifer-, Acrospirifer-, Euryspirifer-, Paraspirifer-, and Multiplicatispirifer-like morphotypes, which are briefly introduced. The new systematics is characterized by different clades, the European/North African delthyridoid spiriferid clade, the North American delthyridoid spiriferid clade, the Asian delthyridoid spiriferid clade, the Malvinokaffric delthyridoid spiriferid clade, and the delthyridoid multiplicated spiriferid clade. Each of them is described in a cladistic and in a phylogenetic way. Their phylogenetic relationship sheds new light on palaeobiogeographical interpretations for the different stages from Late Silurian to early Middle Devonian time. A tendency for increasing endemicity is seen until the end of the Early Emsian, which is interrupted by short term regional faunal exchange within a province or within a realm, followed by a loss of endemicity resulting in global distribution of brachiopod genera until the end of Givetian time. The Old World Realm is re-defined due to the lack of phylogenetic relationship between its faunas and subdivided into the European Realm, consisting of the Gondwanan and Avalonian provinces, and the Asian Realm, consisting of the Siberian, Sino, and Mongolian provinces. A reconstruction of Lower Devonian palaeobiographical map is introduced.Delthyridoide Spiriferiden zeichnen sich durch eine weltweite Verbreitung und schnelle Evolution im Paläozoikum aus und gelten als wichtige Werkzeuge für Biostratigraphie und Paläobiogeographie. Es werden Brachiopoden von unterschiedlichen paläobiogeographischen Einheiten direkt miteinander verglichen, um ihre phylogenetische Verwandtschaft zu erforschen und die Paläobiogeographie während des späten Silurs bis zum Eifelium zu verbessern. Die daraus resultierende neue Systematik beruht im wesentlichen auf direktem Vergleich von einzel- und doppelklappigen Schalenmaterial, Innen- und Außenabdrücken sowie Latexabgüssen und Serienschliffen. Die durchgeführten computergestützten kladistischen Analysen haben sich als unzureichend und oft als falsch erwiesen, was durch die infolge der unterschiedlichen Erhaltungsweisen und oft unzureichendem Material erhaltene unvollständige Matrix zu begründen ist. Ein weiteres Problem liegt in der konvergenten Entwicklung der Spiriferiden. Die meisten Merkmale werden unabhängig voneinander in mehreren Entwicklungslinien zu unterschiedlichen Zeitpunkten entwickelt, so dass sich oft auch charakteristische Autapomorphien eines Zweiges nicht erkennen lassen. Die einzelnen Familien und Gattungen lassen sich daher nur in einer Kombination unterschiedlicher Merkmale fassen, anstatt an einer oder wenigen Autapomorphien. Als neue Methode für die taxomische Bearbeitung wird die 3d-Rekonstruktion nach Serienschliffen vorgestellt, die es ermöglicht, Steinkerne und doppelklappiges Schalenmaterial direkt miteinander zu vergleichen. Variationsstatistische Untersuchungen sind, sofern ausreichendes Material vorliegt, an neu aufgestellten Taxa durchgeführt worden, allerdings werden diese aufgrund häufiger unvollständiger Erhaltung und/oder tektonischer Deformation deskriptiv und nicht als taxonomisch entscheidend betrachtet. Brachiopoden, insbesondere Typusmaterial, aus Sammlungen unterschiedlicher Institutionen und Museum sowie eigenes, wenn möglich von der Typuslokalität, aufgesammeltes Material, sind untersucht undrevidierte Diagnosen vom Familien- bis Artniveau aufgestellt worden. Während der Erstellung dieser Arbeit sind 7 neue Familien: Australospiriferidae, Murchisonispiriferidae, Orientospiriferidae, Otospiriferidae, Patriaspiriferidae, Rostrospiriferidae und Trigonospiriferidae; 6 neue Gattungen, davon 1 in offener Nomenklatur: Cyclopterospirifer, Hallispirifer, Parlinispirifer, Murchisonispirifer, Shujiapingensispirifer und gen. nov. B; und 3 neue Arten: Patriaspirifer merriami, Patriaspirifer johnsoni und Murchisonispirifer feldmani aufgestellt worden sowie 1 Taxon als nomen novum: Orientospirifer nakaolingensis wani. Während dieses Projekt sind bereits einige Taxa aufgestellt und publiziert worden, die ebenfalls beschrieben werden: 2 neue Familien: Filispiriferidae and Multispiriferidae; 1 neue Unterfamilie: Multiplicatispiriferinae; 6 Gattungen, davon 1 in offener Nomenklatur: Frequentispirifer, Leonispirifer, Multiplicatispirifer, Ovetensispirifer, Turcispirifer und Genus A; und 9 neue Arten, davon 3 in offener Nomenklatur: Filispirifer hamadae, Leonispirifer leonensis, Multiplicatispirifer foumzguidensis, Oventensispirifer novascotianus, Quiringites arensentiae, Turcispirifer turciae, Multiplicatispirifer cf. foumzguidensis, Quiringites cf. arensentiae und ?Turcispirifer sp. A. Die Brachiopodenfaunen bestehen aus äußerlich sehr ähnlichen Spiriferen, die 6 verschiedene Morphotypen darstellen: Howellella-, Arduspirifer-, Acrospirifer-, Euryspirifer-, Paraspirifer- und Multiplicatispirifer-Morphotypen. Die neue Systematik zeichnet sich durch verschiedene Evolutionszweige aus, den Europäisch/Nordafrikanischen, dem Nordamerikanischen, dem Asiatischen, dem Malvinokaffrischen und dem multiplicaten delthyridoidem Spiriferidenzweig, die jeweils in kladistischer und phylogenetischer Weise beschrieben werden. Die gewonnenen Erkenntnisse über die Verwandtschaftsbeziehungen der einzelnen Faunen führen zu einer Neuinterpretation der Kontinentverteilung und Neudefinition der Faunenreiche innerhalb der einzelnen Stufen des untersuchten Zeitintervalls. Es ist eine zunehmende Provinzialität vom Silur bis zum Ende des frühen Emsiums zu beobachten, die durch zeitweiligen regionalen Faunenaustausch unterbrochen wird. Seit dem späten Emsium setzt Schritt-für-Schritt ein Faunenaustausch ein, der zu einer globalen Verbreitung von Brachiopoden-Gattungen im oberen Givet führt. Das Altweltliche Faunenreich wird in ein Europäisches Faunenreich, bestehend aus einer Gondwana und einer Avalonia Provinz, und in ein Asiatisches Faunenreich, bestehend aus einer Sibirischen, einer Sino und einer Mongolischen Provinz. Eine neue paläobiogeographische Karte für das frühe Devon, die den Südchinesischen Kontinent näher an den Sibirischen Kontinent stellt, wird vorgestellt

Supplemental Data 1

Data set of images of grinding surfaces of Athyris africana n. sp. SMF 94007.2 (right), SMF 94007.3 (center), SMF 94007.4 (left and used for reconstruction)