The first tunicate with a calcareous exoskeleton (Upper Triassic,northern Italy)

The first tunicates with a calcareous exoskeleton are reported from Late Triassic buildup‐slope deposits of the Dolomites. Although examples of this group have been known since the early 1900s from the middle–upper Permian of eastern Asia and Sicily as Khmeria, they were erroneously attributed to rugose corals. These early representatives are small, double‐valved, conical skeletons, which evolved into multi‐plated capsules with up to 35 opercula. The latter are joined along zigzag margins, which in life could probably be opened for the atrial and branchial siphons. The construction and shape of these skeletons distinguish them from plants or other invertebrate phyla, while they share several similarities with living tunicates, specifically to sessile ascidians. Apart from a soft‐bodied genus from the lower Cambrian of China, ascidians are known only from isolated spicules, which occur sporadically from the Lower Jurassic onwards. The calcareous skeleton of these Late Triassic tunicates consists of aragonitic fibres, which form spherulitic or clinogonal microstructures. It seems that the stellate aragonitic spicules of Jurassic to Recent ascidians are a vestige of Permian–Triassic ancestors, which after the Carnian lost the ability to construct compound solid skeletons but partly still retain a soft double‐valved or multi‐operculate cellulose‐like tunic. The following taxa are described as new: Order Khmeriamorpha with the genera Khmeria Mansuy and Zardinisoma gen. nov., and the following species: Khmeria stolonifera (late Permian), Khmeria minima (Late Triassic), Zardinisoma japonicum (late Permian), Z. cassianum, Z. pyriforme, Z. polyplacophorum and Z. pauciplacophorum (all Late Triassic).     

Composition,Seasonal Dynamics,and Long-Term Fluctuations in the Density of Pelagic Polychaetes in Amurskii Bay,Sea of Japan

This is the first survey of the composition and seasonal density dynamics of the pelagic polychaete larvae in Amurskii Bay. This is a summary of the results of two years of observations (1996–1998). The larvae of 14 polychaete families have been recorded. The greatest number of species have been found in the family Spionidae. The most common were the larvae of Harmothoe imbricata (family Polynoidae) and Paraprionospio sp. (family Spionidae). The larvae of polychaetes have been found in plankton throughout the year and have two density peaks. The first peak was due to the spawning of H. imbricata; the second one was provided by representatives of several species of the family Spionidae, mostly Paraprionospio sp., Polydora ciliata, and P. caeca.     

Host‐specific acrothoracid barnacles on Middle Devonian platyceratid gastropods

Borings, attributed to acrothoracic barnacles, occur on the platyceratid gastropod Naticonema lineatum (Conrad) from the Middle Devonian Hamilton Group of western New York and rarely in specimens as old as the Early Devonian. These latter are the oldest known acrothoracid borings are in the fossil record. The borings are consistently developed as laterally compressed, inequilateral pouches exclusively on these gastropods, commonly as dense infestations.

Naticonema shells yielding borings typically occur associated with partially articulated qrinoid remains, and they are sometimes found attached to crinoids in a manner similar to coprophagous Platyceras. In addition to barnacle borings, Naticonema shells often bear thin encrustations of bryozoans which are usually perforated by these borings but sometimes also overgrow them. Barnacles bored live hosts; gastropods prevented shell penetration by producing cyst‐like secondary secretions of calcite beneath acrothoracid boreholes.

The relative antiquity of these borings and their association with coprophagous platyceratids makes their discovery particularly significant in revealing aspects of the early ecology of barnacles. Attachment to the host commensal gastropods was one of the first successful life modes of these crustaceans prior to their later diversification to other habitats. Mississippian and Pennsylvanian occurrences of similarly bored gastropods demonstrate continuity of the barnacle‐gastropod‐crinoid ecological association from the Middle to Late Paleozoic.     

Geological history and phylogeny of Chelicerata

Chelicerata probably appeared during the Cambrian period. Their precise origins remain unclear, but may lie among the so-called great appendage arthropods. By the late Cambrian there is evidence for both Pycnogonida and Euchelicerata. Relationships between the principal euchelicerate lineages are unresolved, but Xiphosura, Eurypterida and Chasmataspidida (the last two extinct), are all known as body fossils from the Ordovician. The fourth group, Arachnida, was found monophyletic in most recent studies. Arachnids are known unequivocally from the Silurian (a putative Ordovician mite remains controversial), and the balance of evidence favours a common, terrestrial ancestor. Recent work recognises four principal arachnid clades: Stethostomata, Haplocnemata, Acaromorpha and Pantetrapulmonata, of which the pantetrapulmonates (spiders and their relatives) are probably the most robust grouping. Stethostomata includes Scorpiones (Silurian–Recent) and Opiliones (Devonian–Recent), while Haplocnemata includes Pseudoscorpiones (Devonian–Recent) and Solifugae (Carboniferous–Recent). Recent works increasingly favour diphyletic mite origins, whereby Acaromorpha comprises Actinotrichida (Devonian–Recent), Anactinotrichida (Cretaceous–Recent) and Ricinulei (Carboniferous–Recent). The positions of the Phalangiotarbida (Devonian–Permian) and Palpigradi (Neogene–Recent) are poorly resolved. Finally, Pantetrapulmonata includes the following groups (listed here in their most widely recovered phylogenetic sequence): Trigonotarbida (Silurian–Permian), Uraraneida (Devonian–Permian), Araneae (Carboniferous–Recent), Haptopoda (Carboniferous), Amblypygi (?Devonian–Recent), Thelyphonida (Carboniferous–Recent) and Schizomida (Paleogene–Recent).     

Feeding of the Cymatiid Gastropod, Argobuccinum argus, in Relation to the Structure of the Proboscis and Secretions of the Proboscis Gland

Several gastropod families belonging to the superfamily Tonnaceaare known to bore into the calcareous shells of echinoids andbivalves, but little work has been done on the family Cymatiidae.New observations show that the cymatiid, Argobuccinum argus,which feeds on the tubicolous polychaete, Gunnarea capensis,possesses large proboscis glands which secrete 0.4 to 0.5N H₂SO₄;further, the secretion dissolves greater amounts of pure CaCO₃than does O.47N H₂SO₄, alone. A toxin is also present, whichabolishes responses to light and touch in several invertebrates. The proboscis is atypically pleurembolic, having a permanentproboscis sheath. Retraction is due to the muscles ot the probosciswall, and the contracted proboscis is not inverted. Unlike othermesogastropods, the Tonnacea have three pairs ofglands associatedwith the foregut: proboscis glands, true salivary glands, anda small, partly paired gland of unknown function immediatelyposterior to the buccal mass. It is concluded that because of an abundance of readily availablefood in the form of Gunnarea, A. argus does not drillinto animalsprotected by a calcareous shell. The presence of a CaCO₃ dissolvingmechanism indicates that the Cymatiidae ingeneral may be ableto employ this method of feeding.     

Bioerosion in Late Permian Rugosa from reefal blocks (Hawasina Complex,Oman Mountains): implications for reef degradation

Summary Rugose corals are known from allochthonous Late Permian reefal blocks of the A1 Jil and Ba’id Formation (Hawasina Complex), Oman Mountains. In contrast to many Late Permian Rugosa found elsewhere in the Tethys, they occurred in sponge reefs and contributed to reef construction. The waagenophyllid warm water coral fauna is moderately diverse comprising cerioid, thamnasterioid, and fasciculate taxa. In contrast to sponges, chaetetids, and low-growing reefbuilders, the corals secreted diagenetically stable, most probably Mg-calcitic skeletons. Borings in coral skeletons are consequently well preserved providing important data for the interpretation of reef destructive processes. Thin-section analysis revealed three taxa of infaunal borers includingEntobia Bronn 1837, uncertain thallophyte borings, and borings of unknown bioeroders. Macroborers were more important than microborers, because of the dominance of clionid sponges. Good evidence exists also for the occurrence of two types of undetermined grazers which destroyed the coral surfaces. The amount and distribution of bioerosions is variable among different coral taxa. The fasciculate coralPraewentzelella regulare Flügel 1995 was the favorate substrate. Up to 33% of the calices were bored. Dendroid and compound corals were bored subordinately. Bioerosion of these colonies does not exceed 2%. There is good evidence for substrate preference amongst the borers. Major controlling factors affecting borer distribution are believed to be variations of skeletal density and gross morphology. The borer assemblage could not limit reef accretion significantly. Factors controlling boring activity might have been quality of substrate, sedimentation rate, rapid incrustation of substrates, and competition for food with reef constructors including sponges, chaetetids, and rugose corals.     

Ultrastructure of the extensively developed nuchal organs of Laonice bahusiensis (Annelida: Canalipalpata: Spionidae)

The nuchal organs of annelid Laonice bahusiensis (Spionidae) from northern Europe have been studied using scanning and transmission electron microscopy. L. bahusiensis is the first spionid species in which extensively developed, continuous nuchal organs are described. The nuchal organs of this genus are the longest known among polychaete annelids. They consist of paired double bands extending from the prostomium on a mid‐dorsal caruncle for about 24–30 setigers. Their microanatomy corresponds to the general structural plan of nuchal organs: there are ciliated supporting cells and bipolar sensory cells with sensory cilia traversing an olfactory chamber. The organs are overlaid by a secondary paving‐stone‐like cover and innervated by one pair of longitudinally elongated nuchal nerves. These findings clearly favor the hypothesis that the paired, extensively developed ciliated structures found in some Spionidae are homologous with the prostomial nuchal organs characteristic of polychaete annelids. J. Morphol. 2010. © 2009 Wiley‐Liss, Inc.     

A pyritized polychaete from the Devonian of Ontario

A polychaete from the Middle Devonian Arkona Shale at Hungry Hollow, Arkona, Ontario is preserved in three dimensions in pyrite. The prostomium bears a single median antenna, a pair of lateral antennae and a pair of ventral palps. It is assumed to be fused to a reduced peristomium. The anteriormost three pairs of trunk appendages are modified as tentacular cirri, the third long and biramous. The remainder of the finely annulated trunk bears at least 21 similar biramous parapodia, some of which preserve evidence of chaetae. The postsegmental pygidium is very small and may bear up to two pairs of cirri. The polychaete, Arkonips topororum, falls within the Palpata, Aciculata, among the crown group Phyllodocida. Its remarkable preservation highlights the potential of the Arkona Shale to yield other examples of soft-tissue preservation.     

Endolith associations and their relation to facies distribution in the Middle Devonian of New York State, U.S.A.

Vogel, Klaus, Golubic, Stjepko & Brett, Carlton E. 1987 07 15: Endolith associations and their relation to facies distribution in the Middle Devonian of New York State, U.S.A.
Microborings of endolithic organisms (e.g. algae, fungi) provide valuable indicators of paleoenvironments. The present study documents the distribution of microborings in fossil skeletal substrates from a spectrum of shelf to basinal facies in the Middle Devonian of New York. Endolithic boring in brachiopod shells and corals were prepared using special embedding-casting method and were examined and photographed with SEM. About 13 distinct types of microborings were discovered, presumably produced by microorganisms (pro- and eukaryotic), sponges, bryozoans and worms. Brachiopod holdfast etchings are reported for the first time from Paleozoic rocks. Microborings are non-randomly distributed among Middle Devonian facies. The highest diversity and abundance of borings occurs in diverse brachiopod and coral-bearing calcareous mudstones; which appear to represent shallow, well-aerated and low turbidity environments. Microboring diversity and frequency decreases markedly into mudstones and siltstones recording higher turbidity as well as into dark gray to black basinal shale facies. 'Leiorhynchus' brachiopods from probable deeper water, dysaerobic environments display very low diversity of micro-boring assemblages. They include brachiopod pedicle borings and one endolith which shows similarities to a modem cyanobacterium. If the latter is correctly identified, it brackets all of the studied facies within the photic zone.     

Simulations of the Ontogeny of Spongiophyton, a Devonian Plant

Mathematical representations of the cellular organization anddimensions of Spongiophyton Krusel, a Middle Devonian thallophytewere projected by means of a computer to simulate patterns ofdevelopment and organization. Extrapolation of the cellularpatterns observed on the surface of the fossil may be comparedwith those derived from living plants of which the ontogenycan be directly observed. Spongiophyton is compared in thisrespect with the growth of Protosalvinia (an Upper Devonianplant of enigmatic affinity), Pellia (a thallose liverwort)and Cutleria (a brown alga). The growth pattern of Spongiophytondeveloped by computer shows a closer similarity to that of thepseudoparenchymatous alga Cutleria than to the truly parenchymatousPellia or the fossil Protosalvinia. Computer simulations ofthe growth process throw light on the affinity of Spongiophytonwhich cannot be derived from direct observation of the fossil.Broader applications of computer simulations of tissue organizationand gross morphology are suggested with regard to the studyof living and fossil plants.     

Borings in Quartzite Surf Boulders from the Upper Cambrian Basal Deadwood Formation,Black Hills of South Dakota

Shallow, semi-spherical borings occur in clusters with densities of 1–3.5/cm² in quartzite boulders and in vein quartz from the localized basal conglomerate of the Cambrian-Ordovician Deadwood Formation in the east-central Black Hills of South Dakota. Some borings are superimposed on primary but enigmatic semi-circular structures, 2.5–5 cm in diameter, which are soft-sediment trace fossils formed prior to lithification. The macroborings are the first to be recorded from quartzite and vein quartz. Host boulders were eroded from near vertically dipping Paleoproterozoic quartzites from several different stratigraphic units. The thin boulder conglomerate grades were abruptly changed to sandstone through a layer ≤2 m thick in the Marjuman transgression (regionally correlated to the Cedarina dakotaensis trilobite zone). This transgression occurred prior to the start of the globally recognized Upper Cambrian Paibian Stage. Physically similar rocky-shore settings are widely known from quartzite islands of Cambrian age in Wisconsin, Middle Ordovician age on Ontario's Manitoulin Island, Ordovician-Silurian age in Manitoba, and Devonian age in western Australia. Erosion of quartzite surf boulders of equal or larger size occurred in all those regions, but the Black Hills of South Dakota is the only region where borings in quartzite are documented.     

Agglutinated loricae of some Baltic and Antarctic Tintinnina species (Ciliophora)

The abundance and species composition of diatoms adhering tothe loricae of four agglutinated Tintinnina species, Laackmanniellanaviculaefera, Codonellopsis gaussi, Cd.balechi and Tintinnopsislobiancoi, were determined. Diatoms from the Fragdariopsis group,F.cylindrus and F.pseudonana, dominated on tintinnid loricaefrom the Antarctic waters, whilst Thalassiosira spp. were predominanton loricae from the Baltic Sea. Although tintinnids utilizeddiatoms in the environment, it is not a rule that they use onlythese which are dominant. Our results suggest that certain diatomsare actively selected and agglutinated by particular tintinnidspecies.     

Reassessment of the Devonian problematicum Protonympha as another post‐Ediacaran vendobiont

Protonympha is an enigmatic fossil represented by two species from the Middle Devonian (Protonympha transversa) and Late Devonian (Protonympha salicifolia) of New York. Although interpreted in the past as a polychaete worm or starfish arm, Protonympha is not found with marine fossils, but with fossil plants. This fossil plant community was a swamp woodland of Lepidosigillaria whitei, with ground cover of Haskinsia colophylla, fringing brackish to freshwater coastal lagoons of the Catskill Delta. Protonympha shares with Ediacaran Vendobionta a quilted body of unskeletonized biopolymer that is unusually resistant to burial compaction. In overall form, Protonympha is most like the Ediacaran genus Spriggina. Protonympha has branching and tapering tubular structures radiating from the bottom. These rhizine‐like structures, thallus stratification and internal chambers revealed by petrographic thin sections suggest affinities with lichenized fungi. As for Cambrian Swartpuntia and Ordovician–Silurian Rutgersella, Protonympha may have been a post‐Ediacaran vendobiont.     

Problems in polychaete systematics

Some of the intriguing issues in current polychaete systematics are reviewed. (1) The root of the `polychaete' tree. Currently there are two major hypotheses concerning the root position among polychaetes. One is based on rooting cladograms with outgroups such as Mollusca and result in simple-bodied taxa such as Opheliidae and Questidae forming a basal annelid grade along with Clitellata. Other hypotheses do not use outgroup rooting but involve scenarios on the evolution of the group and would place taxa in Aciculata as basal annelids, thus making Aciculata and Phyllodocida paraphyletic. Molecular sequence data has been of little help in resolving this issue thus far, largely due to limited taxon sampling. (2) Paraphyly. Owing, in part, to a tradition involving the emphasis on differences among taxa, and the application of Linnean ranks (e.g., family), paraphyly is undoubtedly a widespread phenomenon in polychaete systematics. An example of this has been proposed already for Spionidae. If the tree topology and rooting used by Blake & Arnofsky (1999) is correct, Spionidae is made paraphyletic by the recognition of the following four family-ranked taxa; Trochochaetidae, Poecilochaetidae, Longosomatidae and Uncispionidae. Another possible example is seen with Cirratulidae. A preliminary cladistic analysis shows that it is entirely possible that seven other taxa recognised as families may be nested within Cirratulidae. These include Acrocirridae, Ctenodrilidae, Fauveliopsidae, Flabelligeridae, Flotidae, Poeobiidae and Sternaspidae. (3) Problematic taxa. Apart from the problems exposed by the analysis of Cirratuliformia, the position of some of these groups, such as Aberranta, Alciopidae, Hesionides,Lopadorhynchidae, Microphthalmus,Nerillidae, Spinther,Tomopteridae and Sabellariidae, is discussed.     

Bioerosive structures from Miocene marine mobile‐substrate communities in southern Spain,and description of a new sponge boring

Abstract: Neogene palaeoshore sediments are abundantly represented along the Mediterranean coast of Iberia. An outcrop north of the Sierra Tejeda, named La Resinera, exposes concentrations of pebbles and boulders of marble, comprising an upper Miocene marine beach deposit. The high diversity of bioerosion trace fossils present in these boulders includes structures produced by polychaete annelids, demosponges, echinoids and endolithic bivalves, which indicate a shallow shoreface environment. The ichnotaxa represented are Maeandropolydora sulcans, Caulostrepsis taeniola, Entobia geometrica, Entobia ovula, Circolites kotoncensis, Gastrochaenolites torpedo, Gastrochaenolites lapidicus, Gastrochaenolites ornatus and Gastrochaenolites turbinatus. The borings are Tortonian (late Miocene) in age. Also present, and particularly abundant, are large sponge borings that have a single chamber from which radiating canals emerge. This trace fossil is designated as Entobia resinensis isp. nov.     

High diversity and pan-oceanic distribution of deep-sea polychaetes: Prionospio and Aurospio (Annelida: Spionidae) in the Atlantic and Pacific Ocean

Organisms Diversity & Evolution - Prionospio Malmgren 1867 and Aurospio Maciolek 1981 (Annelida: Spionidae) are polychaete genera commonly found in the deep sea. Both genera belong to the...     

Polychaete community structure of the northwestern coast of Mexico: patterns of abundance and distribution

In August 1998, 72 benthic stations were sampled off the northwestern coast of Mexico. A baseline survey of the polychaete macrofauna was carried out for the southernmost portion of the Southern California Bight (SCB) region. Spatial patterns of polychaete assemblages were characterized to determine evidence of environmental impact from point sources of wastewater discharge near the study area, and to define a standard against which to detect future change. A total of 4231 polychaetous annelids representing 38 families and 210 species were identified, which constituted 53% of the total macrofauna. Families with highest species richness were Spionidae (18 spp.), Onuphidae (15 spp.), and Terebellidae (14 spp.). Species with greatest abundances were Spiophanes duplex (490 ind.), Spiophanes fimbriata (330 ind.), Lanassa sp. D (209 ind.), Pectinaria californiensis (138 ind.), and Euclymeninae sp. A (124 ind.). Based on polychaete assemblages, communities for the southernmost SCB were found to be composed primarily of non-opportunistic species, and no numerically dominant species were found. High values of polychaete diversity (H) indicated the majority of the area of study could be considered an undisturbed environment. Principal Components Analysis (PCA) applied to environmental and faunistic parameters indicated depth, organic carbon, and sediment grain size as the main influences determining the polychaete faunal communities.     

Global diversity of polychaetes (Polychaeta; Annelida) in freshwater

A literature review of Polychaeta (Annelida) including Aphanoneura (the oligochaete-like Aeolosomatidae and Potamodrilidae), living in freshwater yielded 168 species, 70 genera and 24 families representing all of the major polychaete clades, but less than 2% of all species. The best-represented families were, in order, Nereididae, Aeolosomatidae, Sabellidae, Spionidae and Histriobdellidae. Fourteen families were represented by a single species and genus. Regions supporting the highest diversity of freshwater polychaetes were in order, Palaearctic, Neotropical, Oriental, Nearctic, Australasian, and Afrotropical. More than half of all species and genera inhabitat lakes and rivers, followed by lagoons/estuaries, which have a high proportion of euryhaline species, and inland seas. Less common, atypical polychaete habitats include subterranean waters, the hyporheic zone of rivers and plant container habitats (phytotelmata). At least three distinct ecological/historical processes appear to account for the colonisation of continental waters: invasion of a clade prior to the break-up of Gondwana, as in Aphanoneura, Namanereis, Stratiodrilus, and Caobangia; relatively recent stranding of individual species (relicts); and the temporary visitation of euryhaline species. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

A New Ichnospecies of Gastrochaenolites Leymerie from the Pleistocene Port Morant Formation of Southeast Jamaica and the Taphonomy of Calcareous Linings in Clavate Borings

The ichnospecies Gastrochaenolites pickerilli isp. nov. is based on ten borings found in a shell of the gastropod Strombus gigas Linné from the Pleistocene (Sangamonian) Port Morant Formation of southeast Jamaica. These borings bear morphological similarities to Gastrochaenolites torpedo Kelly and Bromley but differ from all other Gastrochaenolites ispp. in having prominent and numerous calcareous meniscate structures arrayed adjacent to one side of the boring. These menisci are concave towards the center of the boring and are the remnants of calcareous tubes that lined earlier boreholes, that the boring bivalve treated as part of the lithified substrate when relocating. They are thus evidence of the former positions of borings that, unusually, were breached as the bivalve migrated sideways. Although this was a common behavior for Gastrochaenolites-producing bivalves within this substrate, the reason for it occurring is uncertain.