New data on molluscs and their shell microstructures from the Middle Cambrian Gowers Formation, Australia

Abstract: Numerous new cases of preserved shell microstructure were discovered in molluscs from the Middle Cambrian Gowers Formation (Ptychagnostus atavus/Peronopsis opimus Zone, Floran Stage) in the Georgina Basin, Australia. The new data provide further evidence that, by the Middle Cambrian, molluscan shell microstructures were diverse, and many molluscs had a complex shell with multiple types of shell microstructure. In addition, many new occurrences of laminar microstructures are described herein. For many, the nature of these laminar microstructures is not known, but in three species the microstructure is foliated calcite, and in at least two the microstructure is more likely to have been calcitic semi‐nacre, a type of microstructure known in brachiopods and bryozoans but unknown in modern molluscs. This commonality among these three closely related lophotrochozoans underscores a similar mechanism of biomineralization. Moreover, these observations suggest a prevalence of calcite‐shelled lineages among molluscs from the Middle Cambrian, a time of calcite seas. In addition, the broad occurrence of laminar, nacre‐like microstructures in many of these fossils reveals how widespread these strong (fracture‐resistant) microstructures were in Middle Cambrian molluscs. Additionally, a few specimens of Yochelcionella preserve imprints of a bilaterally symmetrical pair of muscle scars. New taxa described here include Corystos thorntoniensis gen. et sp. nov., Yochelcionella snorkorum sp. nov., Yochelcionella saginata sp. nov., and Anhuiconus? agrenon sp. nov.     

Mobergellans (Problematica) from the Cambrian of Greenland, Siberia and Kazakhstan

Mobergellans are disc-shaped problematic fossils of phosphatic composition bearing prominent muscle attachment structures on the presumed internal surface. They are found exclusively in sediments of Early Cambrian age, mainly from Laurentia, Baltica and Siberia. Type material ofDiscinella brastadi Poulsen, 1932 from North-East Greenland is re-studied and synonomised withDiscinella micans (Billings, 1871). Specimens from Siberia described asD. brastadi are re-assigned toMobergella hexactina n. sp., and specimens from Siberia described asMobergella radiolata Bengtson, 1968 are re-assigned toM. sibirica n. sp.Aktugaia triangula Mis-Sarzhevsky, 1976 from Kazakhstan is redescribed, and its relationship to other mobergellans is discussed.      

Muscle scars, shell form and torsion in Cambrian and Ordovician univalved molluscs

Runnegar, Bruce 1981 12 15: Muscle scars, shell form and torsion in Cambrian and Ordovician univalved molluscs. Lethaia, Vol. 14, pp. 311–322. Oslo. ISSN 0024–1164. Well preserved muscle scars have been discovered on one or more specimens of the early Palaeozoic univalved molluscs Pelagiella, Matherella, Sinuites and ?Bucania. From this information it is argued that the bellerophonts Sinuites and ?Bucania were untorted, that Pelagiella had undergone about 10° of torsion, and that Matherella may have been a fully torted, hyperstrophic descendant of Pelagiella. Other early gastropods such as Aldanella could have been independently derived from Pelagiella and therefore the Class Gastropoda may be at least diphyletic. In general, small dextrally coiled archaeogastropods such as Aldanella and the pelagiellids are the most common asymmetrical univalves of the Early Cambrian, but rare ultradextral (hyperstrophic) forms are also known. Only pelagiellids are known from the Middle Cambrian, and ultradextral forms dominate in the Late Cambrian. As a result, slit-bearing archaeogastropods are not common before the Ordovician. Yuwenia bentleyi gen. et sp. nov. is a new taxon proposed in this work. *Mollusca, Gastropoda, Monoplacophora, anatomy, evolution, Cambrian, Ordovician, Yuwenia n.g.     

A first report of Sphenothallus Hall, 1847 in the Cambrian of Variscan Europe

Ten specimens of two phosphatic fossils have been recently discovered in lower and middle portions of Middle Cambrian Jince Formation in the Czech Republic. They are attributed to the genus Sphenothallus Hall, 1847 and described as two separate species; comparatively small conchs are described as S. kozaki sp. nov., the much larger specimens characterized by its smooth and partly flexible organo-phosphatic walls of shell are determined as ?S. kordulei sp. nov. Sphenothallus is known to range from Cambrian to Permian and accommodates numerous species. However, its Cambrian distribution is considerably restricted. Generally rare specimens have been described from Lower to Middle Cambrian of Laurentia and from the Lower Cambrian of Gondwana and peri-Gondwana. The new record of Sphenothallus from the Jince Biota represents a notable extension of their geographic range.     

STENOTHECOIDA,A PROPOSED NEW CLASS OF CAMBRIAN MOLLUSCA

Cambridium, Bagenovia, and Stenothecoides, composing the Family Cambridiidae, a monotypic superfamily and an order, were in 1960 assigned (although with a query) to the molluscan class Monoplacophora. The basic error of this assignment, according to the author, was the assumption that these specimens are univalves. One specimen from Siberia and a second from Alaska demonstrate that Stenothecoides is bivalved; Bagenovia was first described as a bivalve, but the implication of two valves was ignored. Short internal ridges normal to the shell margin in Cambridium and Stenothecoides, described by Rasetti and Horný, show little resemblance to features of pelecypod shells. These markings are not homologous to paired muscle scars of monoplacophorans. The asymmetric bivalved shell and internal furrows are interpreted as features of class-rank significance; the extinct class Stenothecoida is proposed to accommodate these genera. These animals are most common in Lower Cambrian, but range into Middle Cambrian. They may have been functionally similar to brachiopods, but were unable to compete with those more efficient bivalves.     

Problematic cap-shaped fossils from the Lower Cambrian of North-East Greenland

Three species ofOcruranus Liu, 1979 are described from the Bastion Formation of North-East Greenland, of late Early Cambrian (middle Dyeran of North American usage) age, representing the youngest record of a genus originally described from the earliest Cambrian Meishucunian Stage of China. An accompanying species, tentatively assigned toXianfengella He &Yang, 1982, seems also to be present in South Australia in strata of late Early Cambrian (Botoman of Siberian usage) age, although also this genus was described originally from the Meishucunian.Ocruranus andXianfengella from China have been interpreted as possibly parts of coeloscleritophoran scleritomes, perhaps halkieriids, rather than individual molluscan shells. Their shell form is not typical of helcionelloids which otherwise dominate the Early Cambrian molluscan record, but conclusive evidence of affinity is not forthcoming from the Greenland records. New taxa:Ocruranus septentrionalis n. sp. andOcruranus tunuensis n. sp.      

Revision of Hyolitha from the Ordovician of Estonia

Topotype specimens of the Middle OrdovicianHyolithes acutus Eichwald, 1840, which is the type of the genus that lent its name to a family, order, class, and even phylum according to some, andH. latus Eichwald, 1860 allow that genus and those species to be firmly established on a sound, morphologic basis. In addition, preservation of the types ofHyolithes striatus Eichwald, 1860 is sufficiently good to warrant reassignment toDorsolinevitus Syssoiev, 1958. In contrast, the type ofH. insularis Eichwald, 1860 is incompletely preserved, and this species is not recognizable beyond the type material. The concept of the family Hyolithidae is revised to more closely conform to the morphology ofHyolithes, with authorship herein ascribed toSyssoiev (1958) rather than toNicholson (1872). The stratigraphic distribution of these taxa suggests thatHyolithes as defined herein first appears in the Middle Ordovician, but extends into at least the Lower Devonian, as suggested by two species from the Barrandian region of the Czech Republic. Their geographic distribution further re-enforces the notion of two distinct paleobiogeographic provinces based on hyoliths, a Mediterranean province and Baltic province, with almost no mixing of hyolith faunas during the Ordovician.      

The oldest Jurassic cyathophorid coral (Scleractinia) from siliciclastic environments of the Kachchh Basin, western India

Cyathophora Michelin, 1843, hitherto well known from the Upper Jurassic and Cretaceous, has been found in the Middle Jurassic (Bajocian) of the Kachchh Basin, western India. Eleven specimens ofCyathophora bourgueti (Defrance, 1826) from the Babia Cliff Sandstone member of the Kaladongar Formation, exposed along the northern scarp of the Kala Dongar, Pachchham Island, Kachchh, are described and illustrated as the earliest Jurassic record of the family CyathophoridaeVaughan & Wells, 1943. It is suggested that the monospecific occurrence ofCyathophora bourgueti was controlled by salinity.      

The brachiopod <Emphasis Type="Italic">Eoobolus</Emphasis> from the Early Cambrian Mural Formation (Canadian Rocky Mountains)

The Early Cambrian brachiopod, Eoobolus, is one of the first representatives of the superfamily, Linguloidea, the defining characteristics of which include the classical morphology of oval shells and a pedicle that emerges from between the two valves. The material described here from the Mural Formation (Jasper National Park, Canadian Rocky Mountains) provides well-preserved muscle scars and larval shells that allow a discussion of the muscle system and the larval morphology of Eoobolus. The dorsal larval shell exhibits a morphology similar to other Cambrian linguloids, but also to paterinids, Mickwitzia muralensis, and some rhynchonelliforms. This suggests that there was a lesser degree of disparity among brachiopod larvae in the Cambrian than there is today. The muscle system of Eoobolus is similar to other linguloids, but differs from that of Recent lingulids and discinids by having one or two more pairs of oblique muscles. New data on the distribution of features characteristic of the family Eoobolidae question the validity of this family.     

Periodic shell decollation as an ecology-driven strategy in the early Cambrian Cupitheca

Shell decollation is a growth strategy that has been adopted by a number of invertebrate taxa to offset the metabolic and ecological disadvantages of shell growth. However, little is known about the origin and evolution of this process. We here describe well-preserved specimens of the hyolith Cupitheca decollata sp. nov. preserving the decollation process, from the early Cambrian Yu'anshan Formation (c. 518 Ma) of South China. Based on a large number of specimens collectively representing different developmental stages, we use high-resolution X-ray microtomography and scanning electronic microscopy to reconstruct the process of decollation in this taxon. Cupitheca is among the earliest known small shelly fossils, and thus our discovery confirms that periodic decollation had evolved by the onset of the Cambrian explosion, reflecting the high intensity of the predator–prey arms race in early Cambrian ecosystems. A comparison between the decollation processes of Cupitheca and other shelly invertebrates suggests that periodic decollation and the associated molecular mechanisms of calcium dissolution, uptake, allocation and deposition may have had multiple independent origins.     

First record of the brachiopod Lingulella waptaensis with pedicle from the Middle Cambrian Burgess Shale

Pettersson Stolk, S., Holmer, L. E. and Caron, J ‐B. 2010. First record of the brachiopod Lingulella waptaensis with pedicle from the Middle Cambrian Burgess Shale. —Acta Zoologica (Stockholm) 91 : 150–162 The organophosphatic shells of linguloid brachiopods are a common component of normal Cambrian–Ordovician shelly assemblages. Preservation of linguloid soft‐part anatomy, however, is extremely rare, and restricted to a few species in Lower Cambrian Konservat Lagerstätten. Such remarkable occurrences provide unique insights into the biology and ecology of early linguloids that are not available from the study of shells alone. Based on its shells, Lingulella waptaensis Walcott, was originally described in 1924 from the Middle Cambrian Burgess Shale but despite the widespread occurrence of soft‐part preservation associated with fossils from the same levels, no preserved soft parts have been reported. Lingulella waptaensis is restudied herein based on 396 specimens collected by Royal Ontario Museum field parties from the Greater Phyllopod Bed (Walcott Quarry Shale Member, British Columbia). The new specimens, including three with exceptional preservation of the pedicle, were collected in situ in discrete obrution beds. Census counts show that L. waptaensis is rare but recurrent in the Greater Phyllopod Bed, suggesting that this species might have been generalist. The wrinkled pedicle protruded posteriorly between the valves, was composed of a central coelomic space, and was slender and flexible enough to be tightly folded, suggesting a thin chitinous cuticle and underlying muscular layers. The nearly circular shell and the long, slender and highly flexible pedicle suggest that L. waptaensis lived epifaunally, probably attached to the substrate. Vertical cross‐sections of the shells show that L. waptaensis possessed a virgose secondary layer, which has previously only been known from Devonian to Recent members of the Family Lingulidae.     

The tube-like structures on the juvenile shells of earliest strophomenides and billingsellides as evidence of their life cycles

Possible life cycle of some ancient plectambonitoids (order Strophomenida) from the Middle Ordovician of Russia is reconstructed based on the well-preserved specimens composing the ontogenetic series. Four regions may be distinguished on their shell surface: protegulum, brephic shell, neanic shell and adult shell. The posterior margin of ventral protegulum bears pedicle sheath, which is a tubular outgrowth with a 40-μm-wide aperture at the distal end. The protegulum and brephic shell have common type of microstructure that possibly is spherular; the neanic and adult shells are fibrous. The strophomenide ontogeny possibly was similar to that of recent discinides. The strophomenide life cycle possibly included the planktotrophic juvenile stage; the protegulum and brephic shell were formed in the water column. The aperture of the pedicle sheath was possibly used as an anal opening of the floating juvenile and as an attachment organ during the settlement; at early adult stages, the sheath erased, the anus closed, and the animal started to lie on the ventral valve. The origin of the order Strophomenida and its relative groups is possibly connected with the loss of the pedicle lobe; judging by some strophomenide morphological features, true pedicle was present in the strophomenide ancestors. The tubes on the ventral umbones of strophomenides and billingsellides are not homologous as pedicle sheaths of strophomenides are formed at the planktotrophic swimming stage, and the tubes surrounding the pedicles of billingsellides were formed by deltidial plates of almost adult shell after settling.     

Pumiliornis tessellatus Mayr, 1999 revisited — new data on the osteology and possible phylogenetic affinities of an enigmatic Middle Eocene bird

Although the avian speciesPumiliornis tessellatus Mayr, 1999 is known from two skeletons from the Middle Eocene of Messel in Germany, its phylogenetic affinities remained enigmatic. The new osteological data presented in this study document thatP. tessellatus had an at least semizygodactyl foot, with a very wide basal phalanx of the fourth toe, and lacked an ossified pons supratendineus on the distal tibiotarsus. Compared to the known zygodactyl and semizygodactyl birds, this tiny Middle Eocene species resembles the late Eocene/early Oligocene taxonEocuculus Chandler, 1999. Anew, tentatively referred wing ofEocuculus from the early Oligocene of France is described and compared withPumiliornis.      

Taxonomy of the genusBrimeura (Hyacinthaceae)

A taxonomic reevaluation of two little-knownBrimeura taxa,B. fontqueri (Pau)Speta andB. duvigneaudii (L. Llorens)Rosselló et al., has been made.Brimeura fontqueri, described from the Iberian peninsula, has been put into synonymy ofB. amethystina (L.)Salisb., since it could not be distinguished on morphological, anatomical or cytogenetic grounds.Brimeura duvigneaudii, from the Balearic Islands, is closely related toB. amethystina and has 2n=28 chromosomes. It differs from the latter by its naked bulbs lacking dark cataphylls, and its narrower leaves and whitish corollas. Accessory chromosomes are reported for the first time in the genus. Karyological instability (with chromosome numbers ranging from 2n=28 to 42) is reported for a population ofB. fastigiata (Viv.)Chouard. A key to the recognized taxa ofBrimeura is provided.     

Mallomonas alphaphora (Chrysophyceae), a new species from Western Australia

A new species ofMallomonas, M. alphaphora (Chrysophyceae), was found in freshwater ponds in the Perth region, Western Australia. It is distinguished from other species ofMallomonas by its very distinctive scale and bristle morphology and is placed in a new section,Alphaphorae, of the genusMallomonas. Dedicated to Prof. DrL. Geitler on the occasion of the 90th anniversary of his birthday.     

A late Burdigalian gobiid fish,<Emphasis Type="Italic">Gobius brevis</Emphasis> (<Emphasis Type="SmallCaps">Agassiz</Emphasis>, 1839), in the Upper Hydrobia Beds in the middle Upper Rhine Graben (W-Germany)

Gobius brevis (Agassiz, 1839) was hitherto exclusively based on articulated skeletons. It first appears in the late Early Miocene (late Burdigalian, Karpatian) and is widespread throughout the Middle Miocene. Its previously known zoogeographical distribution was the western Paratethys and the Styrian Basin in Austria. We here report on articulated skeletons ofG. brevis with otoliths in situ from a temporary outcrop at Edenkoben, located in the middle Upper Rhine Graben, Germany. The strata are assigned to the basal part of the Upper Hydrobia Beds. The otoliths in situ are identical with those ofGobius latiformis Reichenbacher, 1992, which is an otolith-based species occurring in the Karpatian and Middle Miocene of the western Paratethys; henceG. latiformis has to be regarded as a younger synonym ofG. brevis (Agassiz, 1839). The skeletons ofG. brevis from Edenkoben differ from other articulated freshwater or brackish water MioceneGobius species in the number of vertebrae, more elongate body shape, and number of fin rays. Also the otoliths ofG. brevis differ in their overall shape from these species. However, the in situ preserved otoliths ofG. brevis in the Edenkoben locality show a more primitive evolutionary stage than the Middle Miocene otoliths ofG. brevis (formerly described asG. latiformis). Thus, the gobiid fishes from Edenkoben belong to the oldest representatives ofG. brevis and must be late Burdigalian in age.Gobius brevis probably migrated into the middle Upper Rhine Graben from the Paratethys area.      

New data on the enigmatic Ocruranus–Eohalobia group of Early Cambrian small skeletal fossils

Abstract: The Ocruranus–Eohalobia group, whose members were variously considered to be brachiopods, bivalves, chitons, tommotiids and coeloscleritophorans, are difficult to classify because of lack of morphological detail and evidence for skeletal reconstruction. New specimens from South China reveal more information about Ocruranus–Eohalobia and allow progress towards deciphering the skeletal reconstruction and phylogenetic affinity of this enigmatic group. Many specimens have a phosphatic inner and outer coat (mould) with empty space in between that resulted from dissolution of the original shell. Moreover, many of the internal moulds show a previously unknown type of shell microstructure that consisted of stacked layers of highly organized, acicular crystallites that radiated from the apex of the shell towards the aperture. The dissolved shell and needle‐like crystals suggest an original calcareous, probably aragonitic, shell mineralogy. A few specimens also show a polygonal texture in regions that suggests the shell had a thin, prismatic inner shell microstructure. Ocruranus and Eohalobia belong to the same skeleton, and we herein synonymize Eohalobia with the older Ocruranus. Moreover, new specimens from Meishucun reveal a third type of shell plate, similar in form and inferred placement to intermediate valves of chitons. Ocruranus is likely a mollusc, and possibly a member of the chiton stem lineage. If so, then the beginning of the known record of chitons would be extended back from late Cambrian (Saukia Zone; Furongian) to early Cambrian (Meishucunian; Series 1).     

Multinuclear stages of the life cycle ofAcetabularia mediterranea

Summary The reproductive stages of the life cycle ofAcetabularia mediterranea have been studied in Feulgen-stained material. Light-microscopical photographs of secondary nuclei, cyst formation, gamete formation, gamete release, zygote formation and early development of the germlings are presented. The time course of cytological events during gamete formation is described. Mitoses are found between 16 and 24 hours after the induction of cyst germination.The author is indebted to Mrs. S.Artelt who helped with the prepration of the specimens and the photographs.     

Structure of shell muscles in the Cambrian gastropod genus Bemella (Gastropoda: Archaeobranchia: Helcionellidae)

Zones with peculiar microornamentation interpreted as muscle scars were found on the internal molds of the Early Cambrian gastropod genus Bemella Missarzhevsky, 1969 (family Helcionellidae). Shell muscles of helcionellids are reconstructed based on the topographic pattern of muscle scars, i.e., the pedal, cephalic, and mantle retractors are recognized. The reconstruction proposed here of the shell musculature corroborates affinity between ancient gastropod and helcionelloid mollusks.