A revision of Holm's late Mid and Late Cambrian hyoliths of Sweden

Type specimens of Hyolithus tenuistriatus Linnarsson show this species to be the second definitive representative of the North American genus Nevadotheca in Scandinavia; H. subcostatus Wallerius is questionably assigned to that genus. Restudy of the types of the hyolithids Hyolithus pennatulus Holm confirms the earlier assignment of that species to Carinolithes Syssoiev and H. araneus Holm is assigned to Lovenedolithes gen. nov. Morphology of the orthothecids Hyolithus ( Orthotheca ) cor Holm, H . ( O .) excavatus Holm and H . ( O .) lineatulus Holm supports placement of the first in Contitheca Syssoiev and the others in Decoritheca Syssoiev. Hyolithus ( Orthotheca ) stylus Holm, type species of Circotheca Syssoiev, is of special significance as the basis for both a family and an order. The morphology of the species suggests that ordinal status is not warranted. Hyolithus obscurus Holm, on which Linevitus Syssoiev is founded, is known only from poorly preserved internal moulds and is unrecognizable.     

Slapylitidae: a new family of hyolithids (Cambrian–?Devonian; Baltica,Laurentia, Gondwana)

Hyoliths are usually preserved as isolated skeletal elements consisting of conch, operculum, and helens. The occurrence of a conch associated with an operculum is ordinarily exceptional, and the co-occurrence of helens with other skeletal parts is a great rarity. The extraordinary finds of hyolithid conchs associated with opercula in situ are relatively abundant in the Cambrian and Ordovician clastic sediments of the Barrandian area in the Czech Republic. The platyclaviculate operculum with clavicles divided by longitudinal walls into channels characterizes members of the newly established family Slapylitidae fam. nov., which includes two genera: Slapylites Marek, 1980 known from the mid-Cambrian of West Gondwana and Baltica and Nevadalites Marek, 1976 documented from the Late Cambrian of Laurentia. To this family most probably belongs also an operculum from the Cambrian Series 2–Series 3 boundary of North Greenland and poorly known material from the Middle Devonian of Bolivia.     

A Revision of Early To Mid Ordovician Hyoliths from Sweden

Hyolith assemblages of early to mid Ordovician age in Sweden include the hyolithids Hyolithus bisulcatus Holm, H. concinnus Holm and H. innotatus Holm, which are confirmed as representatives of Hyolithes Eichwald. Dorsolinevitus Syssoiev is known from D. dispar (Holm), D. textilis (Holm) and D. vomer (Holm), and Carinolithes Syssoiev is represented by C. hospes (Holm) and C. triumvir (Holm). Sulcavitus caelatus (Holm), the type species that forms the basis for a family and even an order according to some, is redescribed; the order is suppressed. Hyolithus cymbium Holm is transferred to Stelterella gen. nov., and Theca crispata (Boll) is now the type species of Crispatella gen. nov. The unusual orthothecid Quadrotheca quadrangularis (Holm) is redescribed; the concept of Trapezotheca aemula (Holm) is emended, and Semielliptotheca dens (Holm) is transferred to Trapezotheca . Semielliptotheca Syssoiev is poorly known from only one species, S. rosmara (Holm), and that genus is now considered to be unrecognizable. Hyolithus exaratus Hadding is identified as closely related to the gastropod Lytospira norvegica Koken, and specimens of Hyolithus sp. nos 11, 12 and 13 Holm and Hyolithus sp. indet. Wiman, remain to be located.     

Homologous shell microstructures in Cambrian hyoliths and molluscs

Hyoliths were among the earliest biomineralizing metazoans in Palaeozoic marine environments. They have been known for two centuries and widely assigned to lophotrochozoans. However, their origin and relationships with modern lophotrochozoan clades have been a longstanding palaeontological controversy. Here, we provide broad microstructural data from hyolith conchs and opercula from the lower Cambrian Xinji Formation of North China, including two hyolithid genera and four orthothecid genera as well as unidentified opercula. Results show that most hyolith conchs contain a distinct aragonitic lamellar layer that is composed of foliated aragonite, except in the orthothecid New taxon 1 that has a crossed foliated lamellar microstructure. Opercula are mostly composed of foliated aragonite and occasionally foliated calcite. These blade or lath‐like microstructural fabrics coincide well with biomineralization of Cambrian molluscs rather than lophophorates, as exemplified by the Cambrian members of the tommotiid‐brachiopod linage. Accordingly, we propose that hyoliths and molluscs might have inherited their biomineralized skeletons from a non‐mineralized or weakly mineralized common ancestor rather than as a result of convergence. Consequently, from the view of biomineralization, the homologous shell microstructures in Cambrian hyoliths and molluscs strongly strengthen the phylogenetic links between the two groups.     

华北寒武系第二统辛集组软舌螺动物化石群*

软舌螺动物是华北板块寒武系小壳化石基本组成之一,也是寒武纪生命大爆发在华北地区的重要见证者。本文系统描述了华北板块南缘-西南缘(河南-陕西地区)寒武系第二统辛集组软舌螺化石9属14种及1未定种。包括直管螺类6属9种:Conotheca australiensis、Cupitheca holocyclata、Cupitheca costellata、Cupitheca? decollata、Tegminites hymenodes、Triplicatella disdoma、Triplicatella xinjia、Protomicrocornus triplicensis和Longxiantheca mira;软舌螺类3属5种及1未定种:Microcornus eximius、Microcornus petilus、Parkula cf. esmeraldina、Parkula bounites、Parakorilithes mammillatus和Parakorilithes sp.。基于大量保存的精美聚合标本,本文准确重建了锥壳和口盖在软舌螺不同属种中的配置。此外,新材料显示华北辛...     

贵州剑河寒武系凯里组脊状单臂螺(Haplophrentis carinatus)埋藏特征及与始海百合共生关系探讨

贵州剑河寒武系凯里组软舌螺化石丰富、埋藏形式多样,其中以脊状单臂螺Haplophrentis carinatus化石数量最多、保存最为完整。过去有关学者对凯里组单臂螺化石的研究主要集中在化石分类学方面,对脊状单臂螺埋藏特征及与其他生物共生关系缺乏深入探讨。本文对324块脊状单臂螺化石标本进行系统研究对比后发现,凯里组脊状单臂螺口盖化石埋藏形式有四类:口盖单独保存、口盖以内模或外模化石形式保存、口盖与锥壳完全绞合保存、口盖与锥壳不完全绞合保存;附肢保存较少;锥壳多以内模化石形式保存;脊状单臂螺内模化石与印痕化石上普遍出现类似软体保存形成的三分叉结构,这类三分叉结构属于后期埋藏因素造成的次生结构。文中还对脊状单臂螺与始海百合共生关系进行探讨,并将二者共生关系归为偏害共生。     

SKELETAL MICROSTRUCTURE OF HELENS, LATERAL SPINES OF HYOLITHIDS

Abstract:  In hyolithids the skeleton consists of four elements: a conch, an operculum and a pair of long, logarithmically curved 'spines' called helens. These last elements are rarely preserved, and have therefore remained poorly known and enigmatic. We have studied three-dimensionally preserved helens of the species ' Hyolithes ' lanceolatus , from the Permian of New South Wales, Australia, and ' Hyolithes ' groenwalli , from the Cambrian of Bornholm, Denmark. Helens were massive calcareous elements. Their original microstructure, herein reported for the first time, consisted of concentric lamellae surrounding a narrow elongated core, which may have been rich in organic matter. This concentric pattern resulted from the successive accretion of shell material at the proximal, internal portion of helens. This growth model is in accordance with helen morphology and with the presence of a characteristic surface sculpture of overlapping lamellae.     

Plywood‐like shell microstructures in hyoliths from the middle Cambrian (Drumian) Gowers Formation,Georgina Basin,Australia

Hyoliths are a group of Palaeozoic fossils with calcareous shells whose affinities remain controversial. As their shells were originally aragonitic, their fossils are usually coarsely recrystallized, and few data on their microstructure are available. We report hyoliths from the middle Cambrian (Drumian, Floran) Gowers Formation of the eastern Georgina Basin, Queensland. These are preserved as phosphatic internal moulds, often with the inner layers of the shell also partly replaced by phosphate. Microstructural details preserved by this early diagenetic phosphatization show that these hyolith conchs were originally composed of fibrous crystallites, c. 0.5 μm wide, parallel to one another and to the inner surface of the shell. In several species, the fibres are arranged in a plywood‐like structure composed of multiple lamellae with a different fibre orientation in each lamella: often they are transversely oriented (relative to the long axis of the conch) in the inner part of the wall and longitudinally oriented in the outer part. Opercula also show a microstructure of parallel fibres. The lamello‐fibrillar microstructure we report from hyoliths is reminiscent of microstructures of many Cambrian molluscs; that this microstructure is found in both conchs and opercula suggests that these structures are serial homologues of one another, and in this respect they resemble brachiopod valves. As with many other biological plywoods, the hyolith shell probably records self‐organization in a liquid‐crystal‐like organic matrix. This provided a straightforward way to construct a material that could resist stresses from different directions, offering an effective defence against predators.     

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.      

Operculum regeneration following failed predation in the Silurian gastropod Oriostoma

Regeneration of the calcareous rigiclaudent operculum following severe, non‐lethal, predatory attacks is described in two specimens of the characteristic Silurian gastropod Oriostoma from the English Midlands. This is the first record of the regeneration of opercula in Palaeozoic gastropods and of opercula conjoined with shells that have been repaired after significant, failed, durophagous attacks. In one specimen, a series of repaired injuries culminates in a repaired aperture closed by the operculum. In the second specimen, new shell growth has failed to recover the full extent of the original broken whorl, and the new aperture and operculum, though well developed, are smaller than the originals. Both opercula are unusually thin centrally when compared to other, strongly domed, Oriostoma opercula.     

Gracilitheca astronauta n. sp. and Nephrotheca sophia n. sp. (Hyolitha,Orthothecida) from the Cambrian of Czech Republic

Two orthothecid hyoliths, Gracilitheca astronauta n. sp. and Nephrotheca sophia n. sp. are described from the “middle” Cambrian Buchava Formation of the Skryje-Tý?ovice Basin in the Czech Republic. The new forms are based on about thirty well-preserved external and internal moulds of both conchs and opercula. Gracilitheca astronauta n. sp. is classified under the family Gracilithecidae Sysoev, 1972 and may be distinguished by clearly triangular cross-section of the conch, whereas Nephrotheca sophia n. sp. is characterized by transversally elongated cardinal processes reaching the lateral opercular margin and is combined with the family Orthothecidae Sysoev, 1958 emended Malinky, 2009. Several specimens of G. astronauta show operculum preserved inside of the conch. Stratigraphic ranges and geographic distributions within the Skryje-Tý?ovice Basin are established for both taxa.     

Biology of the Hyolitha

Hyoliths are Paleozoic fossils that have a calcareous exoskeleton consisting of an elongate, usually bilaterally symmetrical cone, a close fitting operculum, and a pair of curved appendages. Their skeletal ultrastructure resembles the crossed-lamellar shell layers of some molluscs. Several specimens from the Ordovician of France and the Cambrian of Antarctica have parts of the gut preserved by infilling matrix, showing that both mouth ad anus were located near the cone aperture. Muscle scars in other hyolith shells indicate that the animal had a series of dorsoventral and longitudinal, or longitudinal and circular muscles, which operated through a hydrostatic skeleton to protract and retract the head, to open and close the operculum, and to move the appendages. Although the shell form and skeletal ultra-structure of hyoliths are of a molluscan type, the muscle insertions suggest that the hyolith cone is not homologous with the dorsal exoskeleton of primitive molluscs. Hyoliths probably constitute a small extinct branch of phylum size, related to the Mollusca and the Sipunculoidea. All three groups may have had common ancestors in the late Precambrian.     

Operculum ultrastructure in leech cocoons

Clitellate annelids (i.e., oligochaetes including leeches) secrete cocoons as part of their normal reproductive cycle. Typically, the cocoon sheath is passed over the head of the leech and sealed at both ends by opercula (i.e., glue‐like material secreted by the clitellum). Both the fibrous cocoon wall (CW) and opercula are chemically‐related biomaterials that share unusual physiochemical properties, including thermal and chemical resiliency. To explore the underlying morphology of the operculum, we examined cocoons from four leech species (i.e., Myzobdella lugubris, Theromyzon tessulatum, Erpobdella obscura, and Erpobdella punctata) by transmission (TEM) and scanning electron microscopy (SEM). Transmission electron micrographs of all opercula revealed a common, ultrastructural pattern comprising an electron‐dense mosaic of ordered polygons that surrounded interspersed cavities. The long axes of cavities were often oriented directionally, suggesting that operculum material is pliable prior to solidification and distorted as a consequence of cocoon deposition. Concomitantly, the operculum permeates jagged edges of the cocoon sheath sealing the cocoon, which provides a mechanically strong CW/operculum boundary. SEM of leech opercula revealed globular nanoparticles comparable to that observed in bioadhesives from disparate animal phyla (e.g., mussel, barnacle, sea star), suggesting a convergent mechanism of bioadhesion among animals. J. Morphol. 274:940–946, 2013. © 2013 Wiley Periodicals, Inc.     

Hyolith guts in the Cambrian of northern Australia -turning hyolithomorphs upside down

A single specimen of the early Middle Cambrian (Ordian) orthothecimorph hyolith Gudugu-wan hardmani with preserved gut filling has been discovered in the Montejinni Limestone, Wiso Basin, northern Australia. This represents only the eighth such occurrence worldwide. The specimen is unique among orthothecimorphs in retaining the gut filling in combination with the external mould of the operculum and invites revision of the accepted soft-part recon struction in this group, whereby the opercular lateral processes are now recognized to be ventral. Thus orthothecimorph conchs, if horizontal in life, would have been oriented ventral side down, while hyolithomorphs adopted the reverse orientation: horizontal, ventral side up.     

贵州凯里生物群中软舌螺的初步研究

本文描述了采自贵州台江县革东镇八郎中寒武世凯里生物群中的软舌螺化石,计有1种,3相似种和1个未定种。即：Ambrolinevitus cf.ventricosus,Glossolites?sp..Haplophrentis?cf.carinatus,Linevitus opimus,Hyolithes?cf.cariniferus.凯里生物群中软舌螺特点是：1)壳体通常较大,呈宽锥形;2)背、腹壳区分明显,常具中槽或中脊;3)特别是以软舌螺目中的软舌螺科和线带螺科的软舌螺保存数量最多。完全与欧洲、北美相当层位中出现的软舌螺组合面貌相似;并与澄江动物群中的软舌螺在演化特征上是可衔接的。凯里生物群中的软舌螺除具有世界性属种外。还具地区特有的属种。这对研究软舌螺的地理分布、演化进程以及与凯里生物群组合关系有重要意义。     

Thecate stem medusozoans (Cnidaria) from the early Cambrian Chengjiang biota

Cnidarians are phylogenetically located near the base of the ‘tree of animals’, and their early evolution had a profound impact on the rise of bilaterians. However, the early diversity and phylogeny of this ‘lowly’ metazoan clade has hitherto been enigmatic. Fortunately, cnidarian fossils from the early Cambrian could provide key insights into their evolutionary history. Here, based on a scrutiny of the purported hyolith Burithes yunnanensis Hou et al. from the early Cambrian Chengjiang biota in South China, we reveal that this species shows characters distinct from those typical of hyoliths, not least a funnel-shaped gastrovascular system with a single opening, a whorl of tentacles surrounding the mouth, and the lack of an operculum. These characters suggest a great deviation from the original definition of the genus Burithes, and a closer affinity with cnidarians. We therefore reassign the material to a new genus: Palaeoconotuba. Bayesian inference of phylogeny based on new anatomical traits identifies a new clade, including Palaeoconotuba and Cambrorhytium, as a stem group of sessile medusozoan cnidarians that are united by the synapomorphies of developing an organic conical theca and a funnel-like gastrovascular system. This study unveils a stem lineage of medusozoans that evolved a lifelong conical theca in the early Cambrian.     

贵州台江中寒武世的一些软舌螺化石

该文的软舌螺标本采自贵州省台江县八郎村中寒武世地层,计有Haplophrentis?reesei,H.?carinatus,Hyolithes?cf.idahoensis,H.?cf.cariniferus,H.?sp.,Cambrovitus balangensis gen.sp.nov.,Notabititidae gen.et sp.uncertain.。其层位位于梅树村动物群和澄江动物群之上。地层时代与北美布吉斯页岩的时代相当,两地的软舌螺可直接对比。更为可贵的是在方Haplophrentis?carinatus的腹内视标本中发现了直肠和伸缩肌软体印痕。     

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.     

ASCI OF THE PEZIZALES III: THE APICAL APPARATUS OF EUGYMNOHYMENIAL REPRESENTATIVES

Morphological, developmental and cytochemical examinations were made with light and electron microscopy on the apical apparatuses of four eugymnohymenial species, Pyronema domesticum, Ascodesmis sphaerospora, Coprotus winteri and C. lacteus. Ascal tips in all four species were notably thinner walled than the rest of the ascus. Ultrastructurally, demarcation of the opercula was enhanced after staining with silver methenamine. Wide zones of dehiscence are formed in the outer layer of A. sphaerospora and C. winteri. In P. domesticum the outer layer of the operculum is differentially stained from the rest of the ascal wall. Wall dimensions in the multispored asci of C. winteri are approximately three times greater than those in the eight-spored asci of C. lacteus. The shape of the apical apparatus of C. lateus is almost identical to that of C. winteri. Morphological and cytochemical similarities of the apical apparatuses in Ascodesmis, Pyronema and Coprotus help demonstrate greater relationship between these taxa and support the belief that these taxa are most closely related to members of the Otideaceae and Aleuriaceae.     

Comparative morphology of the amphibian opercularis system: I. General design features and functional interpretation

The morphology of the opercularis system of anuran and caudate amphibians suggests that it acts to produce motion of the operculum that in turn produces fluid motion within the inner ear. The operculum and opercularis muscle form a lever system, with a narrow connection between the operculum and otic capsule acting as a fulcrum about which the operculum moves in response to forces applied via the muscle. The opercula of many species possess a muscular process on which the muscle inserts, thereby increasing the moment arm through which the muscle acts. The tonicity of the opercularis muscle allows tensile forces produced by substrate vibration or other mechanical energy applied to the forelimb to be effectively transmitted to the operculum; the elasticity of the connective tissue holding the operculum in place should act to return the operculum to its original position. The opercularis systems of frogs and non-plethodontid salamanders are similar structurally and functionally; that of plethodontid salamanders is structurally distinct but also functions as a lever system. Fluid motion produced by opercular motion could stimulate various end organs of the inner ear; the saccule, lagena, and amphibian papilla are in close approximation and wave energy could directly affect their otoconial or tectorial structures. In those anurans with a tympanic ear, the stapedial footplate and operculum articulate, but this articulation allows both to move independently. The stapes-tympanum complex and opercularis system therefore appear to be independent functional systems, and it is unlikely that the opercularis system modulates middle ear responsiveness. The general design of the opercularis system is consistent with a function in reception of substrate vibrations.