Chondrichthyan‐like scales from the Middle Ordovician of Australia

Abstract: Microvertebrate sampling of the Stairway Sandstone (Darriwilian, Middle Ordovician, central Australia) has yielded scales that are chondrichthyan‐like in their overall construction, and Tantalepis gatehousei gen. et sp. nov. is erected here to describe these specimens. Tantalepis gatehousei gen. et sp. nov. is the stratigraphically oldest microsquamous taxon described thus far, and the ‘shark‐like’ appearance of the scales may extend the chondrichthyan lineage back into the Middle Ordovician. The presence of ‘shark‐like’ scales in the fossil record some 50 myr prior to the first articulated chondrichthyan body fossils and 44 myr before the first clearly identifiable chondrichthyan teeth suggests there is a considerable scope for the recovery of articulated specimens with which to document the early history of crown gnathostomes. Traditional hypotheses of phylogenetic relationships among early jawed vertebrates were recently challenged by the proposal of a radically different tree topology. However, the development of a new data set specifically addressing scale‐based characters is required before taxa such as Tantalepis, that are based upon disarticulated remains alone, can be firmly placed within the emerging, revised, evolutionary narrative.     

Diverse middle Ordovician palaeoscolecidan worms from the Builth‐Llandrindod Inlier of central Wales

Abstract: Palaeoscolecidan worms are rare, Early Palaeozoic fossils with uncertain affinities within the Ecdysozoa. They are locally abundant in the Cambrian and scattered in the Early Ordovician, but very sparse thereafter. Forty‐four specimens have been collected from the Middle Ordovician of the Builth‐Llandrindod Inlier of Mid Wales and include well‐preserved material assigned to seven new genera, with four additional species in open nomenclature. An additional specimen from the Arenig Pontyfenni Formation of South Wales is also described in open nomenclature. The total demonstrates much greater palaeoscolecid diversity than hitherto suspected for this time. The specimens are preserved as cuticle fragments in shales and siltstones, often of submillimetre size but in many cases with excellent preservation. The level of detail preserved in some is equal to that found in Cambrian phosphatized faunas. The new approach to collecting, and the recognition that this material can yield taxonomically useful information, opens new avenues for palaeoscolecidan research in siliciclastic environments. The new taxa are the following: Radnorscolex bwlchi gen. et sp. nov., Aggerscolex murchisoni gen. et sp. nov., Bullascolex inserere gen. et sp. nov., Wernia eximia gen. et sp. nov., Ulexiscolex ormrodi gen. et sp. nov., Pluoscolex linearis gen. et sp. nov. and Loriciscolex cuspidus gen. et sp. nov. The high diversity, and the taxonomic separation from known groups described primarily from Cambrian carbonates, implies that palaeoscolecidans either diversified significantly during the Ordovician or were taxonomically segregated between carbonate and siliciclastic settings. Palaeobiological findings also include confirmation that some palaeoscolecid basal cuticles were solid and others reticulate, plates (and platelets) could form by lateral accretion, plates were in part primarily phosphatic and in part organic and that in at least some groups, platelet secretion occurred external to plate secretion.     

Scale structure of putative chondrichthyan Gladbachus adentatus Heidtke & Krätschmer, 2001 from the Middle Devonian Rheinisches Schiefergebirge,Germany

Gladbachus adentatus is a putative chondrichthyan, known only from the holotype specimen, which comprises an articulated endoskeleton complete from head to pelvic region with the squamation also preserved. The scales superficially resemble those of placoderms more than sharks, in having a similar gross morphology, lamellar cellular bone forming the base and upright dentinous tubercles comprising the crown. The odontocytic mesodentine in the tubercles is comparable to that in the Osteostraci and in some acanthodian taxa, known only from isolated scales, and is probably the plesiomorphic form of dentine for Gnathostomata.     

Early Palaeozoic dentine and patterned scales in the embryonic catshark tail

Regular scale patterning, restricted to the caudalmost tail and organized into two opposing rows on each side of the tail, is observed in few chondrichthyans. These evenly spaced scales, in dorsal and ventral rows, develop in an iterative sequence from the caudal tip, either side of the notochord. They are subsequently lost as a scattered pattern of placoid scales develops on the body and fins. An identical organized pattern is observed in tail scales of Scyliorhinus canicula (catshark), where the expression of sonic hedgehog signal is restricted to the epithelium of developing scales and remains localized to the scale pocket. Regulation of iterative scale position by sonic hedgehog is deeply conserved in vertebrate phylogeny.These scales also reveal an archaic histological structure of a dentine type found in the oldest known shark scales from the Ordovician and Silurian. This combination of regulated pattern and ancient dentine occurs only in the tail, representing the primary scalation. Scattered body scales in elasmobranchs such as S. canicula originate secondarily from differently regulated development, one with typical orthodentine around a central pulp cavity. These observations emphasize the modular nature of chondrichthyan scale development and illustrate previously undetected variation as an atavism in extant chondrichthyan dentine.     

Early scale development in Heterodontus (Heterodontiformes; Chondrichthyes): a novel chondrichthyan scale pattern

In two species of Heterodontus, H. portusjacksoni and H. galeatus, the first scales to develop form two opposing rows along the caudal fin axis on both the left and right sides of the fin. The opposing rows originate from an initial scale located on either side of the posterior tip of the caudal fin, with subsequent scales erupting in a posterior to anterior direction along the tail axis. These scale rows may strengthen tail movements, providing aeration in the egg case, but are lost later in ontogeny. Development of subsequent body scales shows a more irregular origin and arrangement, from anterior to posterior, to cover the dorsal and ventral lobes of the caudal fin. Although the early developmental pattern of the scale associated with the Heterodontus caudal fin has not been previously described, several chondrichthyan taxa, including chimeroids, likewise possess ordered rows of flank scales early in ontogeny that are subsequently lost. These ordered scales contrast with previous suggestions that chondrichthyan scale development is entirely random. Instead, regulated and sequential development of scales may be a plesiomorphic character for both chondrichthyans and osteichthyans, with the less organized arrangement in later ontogenetic stages being a derived condition within Chondrichthyes.     

Early Ordovician bivalves from Dali, West Yunnan, China

A bivalve fauna of Early Ordovician (late Arenig) age is described from the Hsiangyang Formation of the eastern part of West Yunnan, China. The fauna contains elements in common with Early Ordovician faunas of southern Gondwanan areas such as the Montagne Noire and Morocco and with those of Avalonia, together with several previously undescribed taxa. The following taxa are new: Biseriodonta simplex gen. et sp. nov.; Glyptarca sinensis sp. nov.; Trigonoglyptarca magna gen. et sp. nov.; Erhaiconcha xiangyangensis gen. et sp. nov.; Fasciculodonta impressa gen. et sp. nov.; Yunnanoredonia laevis gen. et sp. nov.; Daliella gen. nov.; Goniophorina ( Goniophorina ) contracta sp. nov.; Haidongoconcha radialis gen. et sp. nov.; Eopterinea aequiconcha gen. et sp. nov. The bivalve fauna includes the most diverse glyptarcid fauna and the earliest nuculanid and paracyclid known hitherto.   Eastern West Yunnan belonged to the Indochina terrane and it is concluded that the bivalve fauna represents a high-latitude assemblage and that the Indochina terrane should be considered a component part of the Peri-Gondwanan continent in the Ordovician; its bivalve faunas contrast with those of neighbouring terranes which have affinities with those of lower latitudes.     

A new paleozoic Symmoriiformes (Chondrichthyes) from the late Carboniferous of Kansas (USA) and cladistic analysis of early chondrichthyans

Background

The relationships of cartilaginous fishes are discussed in the light of well preserved three-dimensional Paleozoic specimens. There is no consensus to date on the interrelationship of Paleozoic chondrichthyans, although three main phylogenetic hypotheses exist in the current literature: 1. the Paleozoic shark-like chondrichthyans, such as the Symmoriiformes, are grouped along with the modern sharks (neoselachians) into a clade which is sister group of holocephalans; 2. the Symmoriiformes are related to holocephalans, whereas the other Paleozoic shark-like chondrichthyans are related to neoselachians; 3. many Paleozoic shark-like chondrichthyans, such as the Symmoriiformes, are stem chondrichthyans, whereas stem and crown holocephalans are sister group to the stem and crown neoselachians in a crown-chondrichthyan clade. This third hypothesis was proposed recently, based mainly on dental characters.

Methodology/Principal Findings

On the basis of two well preserved chondrichthyan neurocrania from the Late Carboniferous of Kansas, USA, we describe here a new species of Symmoriiformes, Kawichthys moodiei gen. et sp. nov., which was investigated by means of computerized X-ray synchrotron microtomography. We present a new phylogenetic analysis based on neurocranial characters, which supports the third hypothesis and corroborates the hypothesis that crown-group chondrichthyans (Holocephali+Neoselachii) form a tightly-knit group within the chondrichthyan total group, by providing additional, non dental characters.

Conclusions/Significance

Our results highlight the importance of new well preserved Paleozoic fossils and new techniques of observation, and suggest that a new look at the synapomorphies of the crown-group chondrichthyans would be worthwhile in terms of understanding the adaptive significance of phylogenetically important characters.     

PERMIAN OSTRACODS FROM THE LERCARA FORMATION (MIDDLE TRIASSIC–CARNIAN?), SICILY,ITALY

Abstract: A rich, diverse Permian ostracod fauna has been recovered from the red and grey, laminated shales and quartz‐rich shales of the Triassic Lercara Formation. Forty‐seven species have been identified, 13 of which are newly described here; they belong to 26 genera of which three are new: Anahuacia lercaraensis sp. nov., Aurigerites siciliaensis sp. nov., Bairdia portellaensis sp. nov., Cristanaria? katyae sp. nov., Fabalicypris gruendeli sp. nov., Lethiersa salomonensis gen. et sp. nov., Lethiersia sinusoventralis gen. et sp. nov., Portella trapezoida gen. et sp. nov., Siciliella elongata gen. et sp. nov., Siciliella infernespinosa gen. et sp. nov., Siciliella prima gen. et sp. nov., Siciliella quadrata gen. et sp. nov., and Siciliella spinorobusta gen. et sp. nov. The assemblages contain or are composed of palaeopsychrospheric forms, which are regarded as index fossils for deep environments. The bathymetry of the different associations in life is evaluated.     

New crinoids from the Baltic region (Estonia): fossil tip‐dating phylogenetics constrains the origin and Ordovician–Silurian diversification of the Flexibilia (Echinodermata)

This study documents previously unknown taxonomic and morphological diversity among early Palaeozoic crinoids. Based on highly complete, well preserved crown material, we describe two new genera from the Ordovician and Silurian of the Baltic region (Estonia) that provide insight into two major features of the geological history of crinoids: the early evolution of the flexible clade during the Great Ordovician Biodiversification Event (GOBE), and their diversification history surrounding the end‐Ordovician mass extinction. The unexpected occurrence of a highly derived sagenocrinid, Tintinnabulicrinus estoniensis gen. et. sp. nov., from Upper Ordovician (lower Katian) rocks of the Baltic palaeocontinent provides high‐resolution temporal, taxonomic and palaeobiogeographical constraints on the origin and early evolution of the Flexibilia. The Silurian (lower Rhuddanian, Llandovery) Paerticrinus arvosus gen. et sp. nov. is the oldest known Silurian crinoid from Baltica and thus provides the earliest Baltic record of crinoids following the aftermath of the end‐Ordovician mass extinction. A Bayesian ‘fossil tip‐dating’ analysis implementing the fossilized birth–death process and a relaxed morphological clock model suggests that flexibles evolved c. 3 million years prior to their oldest fossil record, potentially involving an ancestor–descendant relationship (via ‘budding’ cladogenesis or anagenesis) with the paraphyletic cladid Cupulocrinus. The sagenocrinid subclade rapidly diverged from ‘taxocrinid’ grade crinoids during the final stages of the GOBE, culminating in maximal diversity among Ordovician crinoid faunas on a global scale. Remarkably, diversification patterns indicate little taxonomic turnover among flexibles across the Late Ordovician mass extinction. However, the elimination of closely related clades may have helped pave the way for their subsequent Silurian diversification and increased ecological role in post‐Ordovician Palaeozoic marine communities. This study highlights the significance of studies reporting faunas from undersampled palaeogeographical regions for clade‐based phylogenetic studies and improving estimates of global biodiversity through geological time.     

A new diverse shark fauna from the Wordian (Middle Permian) Khuff Formation in the interior Haushi‐Huqf area,Sultanate of Oman

Abstract: Chondrichthyans are newly reported from the autochthonous Wordian Khuff Formation (middle Permian), cropping out in well‐exposed, low‐palaeolatitude sections in the interior Haushi‐Huqf area of Oman. The shark remains comprise isolated teeth, dermal denticles and fin spines and have been recovered by processing limestone in buffered acetic acid from bulk rock samples. The fauna consists of mainly ctenacanthiform and hybodontiform taxa, identified as Glikmanius cf. myachkovensis, Glikmanius culmenis sp. nov., Omanoselache hendersoni gen. et sp. nov., Omanoselache angiolinii gen. et sp. nov., cf. Omanoselache sp., Reesodus underwoodi gen et sp. nov., Teresodus amplexus gen. et sp. nov., Gunnellodus bellistriatus, Khuffia lenis gen. et sp. nov., Khuffia prolixa gen. et sp. nov. and Euselachii sp. indet. Additional specimens include rare teeth of the lonchidiid cf. ‘Palaeozoic Genus 1’ sp., of the neoselachian Cooleyella cf. fordi and a further indeterminate neoselachian, of an indeterminate petalodont and of the holocephalan Deltodus aff. mercurei and Solenodus cf. crenulatus. Fin spines add a further two taxa, Nemacanthus sp. and Amelacanthus cf. sulcatus, which have neoselachian affinities and therefore an unclear relationship to the recovered teeth. The occurrence of Nemacanthus within this Wordian fauna represents the oldest record of this taxon and its only known occurrence in the Palaeozoic. Of the remaining genera, Glikmanius has previously been recorded from the Wordian, whereas for all the others, this study represents their youngest known stratigraphic occurrence and first occurrence in Guadalupian (middle Permian) strata. This adds significantly to our knowledge of the global diversity of chondrichthyans preceding the end‐Guadalupian biotic crisis. Palaeogeographically, for all taxa, this study represents the first record from the western fringe of the marine Neotethyan basin, and only Cooleyella was previously known from the southern (Gondwanan) part of the Pangaean continental margin.     

Description of a new family,new genus,and two new species of deep‐sea Forcipulatacea (Asteroidea), including the first known sea star from hydrothermal vent habitats

Based on a phylogenetic analysis of undescribed taxa within the Forcipulatacea, a new family of deep‐sea forcipulatacean starfishes, Paulasteriidae fam. nov., is described from deep‐sea settings. Paulasterias tyleri gen. et sp. nov. was observed at recently documented hydrothermal vents on the East Scotia Ridge, Southern Ocean. A second species, Paulasterias mcclaini gen. et sp. nov. was observed in deep‐sea settings in the North Pacific, more distant from hydrothermal vents. Both species are multi‐armed (with between six and eight arms), with a fleshy body wall, and a poorly developed or absent adoral carina. Here, we include discussions of pedicellariae morphology, feeding biology, and classification. © 2015 The Linnean Society of London     

Branching out: a remarkable new branching syllid (Annelida) living in a Petrosia sponge (Porifera: Demospongiae)

We describe the morphology and biology of a previously unknown form of branching annelid, Ramisyllis multicaudata gen. et sp. nov. , an endosymbiont of shallow‐water marine sponges (Petrosia sp., Demospongiae) in northern Australia. It belongs to the polychaete family Syllidae, as does Syllis ramosa McIntosh, 1879, the only other named branching annelid, which was collected from deep‐water hexactinellid sponges during the 1875 Challenger expedition. It differs from S. ramosa in parapodial and chaetal morphology. Ramisyllis multicaudata gen. et sp. nov. has segments of several types, including specialized posterior segments on the emergent portions of the worm, and simplified elongate segments that bridge larger cavities in the sponge interior. Aside from the obvious branching form, the new annelid is similar to Parahaplosyllis, differing from it in lacking pharyngeal armature and in the details of the parapodial chaetae and dorsal cirri. Molecular evidence from 16S and 18S rDNA supports a sister‐group relationship with Parahaplosyllis, with both being sister to Trypanosyllis and Eurysyllis. The phylogenetic position of R. multicaudata gen. et sp. nov. indicates that branching has evolved independently in Ramisyllis gen. nov. and Syllis. This is supported by differences in the branching process between the two taxa: in S. ramosa branching is initiated by segment addition at the parapodium, whereas in R. multicaudata gen. et sp. nov. segments are added from a region between parapodia. A model for branching in R. multicaudata gen. et sp. nov. is proposed and possible developmental processes underlying branching in Annelida, and body symmetry comparisons with other invertebrates, are also discussed. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 481–497.     

Discrimination of extant Pan species and subspecies using the enamel–dentine junction morphology of lower molars

Previous research has demonstrated that species and subspecies of extant chimpanzees and bonobos can be distinguished on the basis of the shape of their molar crowns. Thus, there is potential for fossil taxa, particularly fossil hominins, to be distinguished at similar taxonomic levels using molar crown morphology. Unfortunately, due to occlusal attrition, the original crown morphology is often absent in fossil teeth, and this has limited the amount of shape information used to discriminate hominin molars. The enamel–dentine junction (EDJ) of molar teeth preserves considerable shape information, particularly in regard to the original shape of the crown, and remains present through the early stages of attrition. In this study, we investigate whether the shape of the EDJ of lower first and second molars can distinguish species and subspecies of extant Pan. Micro‐computed tomography was employed to non‐destructively image the EDJ, and geometric morphometric analytical methods were used to compare EDJ shape among samples of Pan paniscus (N = 17), Pan troglodytes troglodytes (N = 13), and Pan troglodytes verus (N = 18). Discriminant analysis indicates that EDJ morphology distinguishes among extant Pan species and subspecies with a high degree of reliability. The morphological differences in EDJ shape among the taxa are subtle and relate to the relative height and position of the dentine horns, the height of the dentine crown, and the shape of the crown base, but their existence supports the inclusion of EDJ shape (particularly those aspects of shape in the vertical dimension) in the systematic analysis of fossil hominin lower molars. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Permian Cynodontia (Theromorpha) of Eastern Europe

A collection of Cynodontia from the Permian of Eastern Europe is revised. The taxonomic position of a number of previously described forms is reconsidered. New taxa, Novocynodon kutorgai gen. et sp. nov. (Thrinaxodontidae) from the Middle Permian and Sludica bulanovi gen. et sp. nov. (Procynosuchidae) from the Upper Permian, are described.     

New planthoppers of the tribe Achilini (Homoptera,Fulgoroidea, Achilidae) from Baltic amber

New taxa of Achilini (Achilidae) are described from Baltic amber: Paratesum rasnitsyni gen. et sp. nov., Protomenocria notata gen. et sp. nov., Psycheona variegata gen. et sp. nov., P. striata sp. nov. Protepiptera kaweckii Usinger, 1939 (= Cixidia christinae Lefebvre, Bourgoin et Nel, 2007, syn. nov.) is redescribed with designation of a neotype. “Cixius” testudinarius Germar et Berendt, 1856, “C.” longirostris Germar et Berendt, 1856 and “Oliarus” oligocenus Cockerell, 1910 are transferred to Achilini. A key to the genera of Achilidae known from Baltic amber is provided.     

A Tremadocian (Early Ordovician) palaeoscolecidan worm from graptolitic shales in Hunan Province,South China

A new locality exposing Tremadocian (Early Ordovician) graptolitic shales in Hunan Province, South China, has yielded an exceptionally well‐preserved annulated worm. This palaeoscolecidan is described as Waflascolex changdensis gen. et sp. nov. and reveals extremely fine detail of the cuticle organization and plating array. The new taxon is characterized by three critical characters: incomplete plate rows that occur only on the posterior end of the worm (except in the posterior‐most area) and do not extend over the entire circumference or along the entire trunk; a regular rhomboidal array of platelets around intercalations; and reduced cuticular organization at the posterior termination. The unique cuticular organization and platelet ornamentation in the new taxon offer insight into functional differentiation of plates in the scleritome. Palaeoscolecid distribution through the early Palaeozoic is reviewed, showing that the worms were widespread in the Cambrian and Ordovician, but became more restricted during the Silurian. Ordovician palaeoscolecidans are diverse in scleritome architecture, and strikingly different from Cambrian taxa, indicating that this group diversified as part of the Great Ordovician Biodiversification Event.     

NEOSELACHIANS FROM THE UPPER CAMPANIAN AND LOWER MAASTRICHTIAN (UPPER CRETACEOUS) OF THE SOUTHERN PYRENEES, NORTHERN SPAIN

Abstract: Bulk sampling of upper Campanian to lower–middle Maastrichtian coastal and lagoonal deposits in five sections of the Tremp Formation in the south‐central Pyrenees yielded numerous neoselachian teeth. The fauna comprises nine taxa of which three species and one genus are new: Hemiscyllium sp., Lamniformes indet., Paratrygonorrhina amblysoda gen. et sp. nov., Coupatezia trempina sp. nov., Coupatezia sp., Coupatezia? sp., Rhombodus ibericus sp. nov. and Igdabatis indicus. The neoselachian fauna is dominated by small nectobenthic rays. This composition resembles assemblages known from the marine Upper Cretaceous, but differs from nearby localities of the Basque‐Cantabrian region and continental selachian associations of the French Pyrenees. The results indicate that Rhombodus might not be a reliable biostratigraphic marker for the Maastrichtian. The faunal composition suggests a shallow trans‐Tethyan connection between Eurasia and India at the end of the Cretaceous Period.     

THE FINE STRUCTURE AND SCALE FORMATION OF CHRYSOLEPIDOMONAS DENDROLEPIDOTA GEN. ET. SP. NOV.(CHRYSOLEPIDOMONADACEAE FAM. NOV., CHRYSOPHYCEAE)1

Chrysolepidomonas gen. nov. is described for single-celled monads with two flagella, a single chloroplast, and distinctive canistrate and dendritic scales. The type species, Chrysolepidomonas dendrolepidota sp. nov., is described for the first time. The canistrate scales bear eight “bumps” on the top surface, and the dendriticscales have a tapered base with a quatrifid tip. These organic scales are formed in the Golgi apparatus and storred in a scale reservoir. The scale reservoir is bounded on two sides by the R₁ and R₂ in microtubular roots of the basal apparatus. The cyst (=stomatocyst, statospore) forms endogenously by means of a silica deposition vesicle. The outer cyst surface is smooth, and the pore region is unornamented. Two other organisms bearing canistrate and dendritic scales, previously assigned to the genus Sphaleromants, are transferred to the genus Chrysolepidomonas. They are C.angalica sp. nov. and C. marine(Pienaar) comb. nov. The distinguishing features of Chrysolepidomonas and Sphaleromantis are discussed. A new family, Chrysolepidomonadceae fam. noc., is described for flagellates covered with organic scales.