Ultrastructure of the Jurassic serpulid tubes-phylogenetic and paleoecological implications.

The ultrastructural diversity of the Middle and Late Jurassic serpulid tubes from the Polish Basin has been investigated. The inspection of 12 taxa representing the two major serpulid clades allowed for the identification of three ultrastructure types-irregularly oriented prismatic structure (IOP), spherulitic prismatic structure (SPHP), and simple prismatic structure (SP). Six of the studied species are single-layered and six species possess two distinct layers. Ultrastructural diversity corresponds to certain serpulid clades. The members of Filograninae have single-layered tube walls composed of possibly plesiomorphic, irregularly oriented prismatic structure (IOP). Two-layered tubes occur solely within the clade Serpulinae, where the external, denser layer is built of either the ordered spherulitic (SPHP) or simple prismatic microstructure (SP), and the internal layer is composed of irregularly oriented prismatic structure (IOP). Apart from phylogenetic signals provided by the tube ultrastructure, it can be used in analyzing paleoecological aspects of tube-dwelling polychaetes. Compared to the more primitive, irregularly oriented microstructures of Filograninae, the regularly oriented microstructures of Serpulinae need a higher level of biological control over biomineralization. The advent of the dense outer protective layer (DOL) in serpulids, as well as the general increase in ultrastructure diversity, was likely a result of the evolutionary importance of the tubes for serpulids. Such diversity of the tube ultrastructural fabrics allowed for maximizing functionality by utilizing a variety of morphogenetic programs. The biomineralization system of serpulids remains more complex compared to other tube-dwelling polychaetes. Physiologically more expensive tube formation allows for mechanical strengthening of the tube by building robust, strongly ornamented tubes and firm attachment to the substrate. Contrary to sabellids, which perform a fugitive strategy, an increased tube durability allows serpulids a competitive advantage over other encrusters.

Bryozoan-cnidarian mutualism triggered a new strategy for greater resource exploitation as early as the Late Silurian.

Bryozoans were common benthic invertebrates in the Silurian seas. The large biodiversity among Silurian benthic organisms prompted diversified interactions, and as a result bryozoans hosted many other organisms as symbionts. Here we analyse the cystoporate bryozoan Fistulipora przhidolensis and unidentified trepostomes intergrown with auloporid tabulate corals and putative hydrozoans. The material comes from the uppermost Prídolí Series (Late Silurian) of the Sõrve Peninsula, Saaremaa, Estonia. Our analysis shows that the interaction was beneficial for both organisms-cnidarians benefited from feeding currents created by the host bryozoan, while the latter benefited from the protection from predators by cnidae, it can thus be classified as mutualism. Such associations are common in modern seas. The analysed organisms are typically encrusting when the symbiosis is absent, when intergrown they display erect, branching morphologies, raised over the substratum, thus exploiting a higher suspension-feeding tier. While similar associations were known from the Devonian, we demonstrate that this novel ecological strategy for greater resource exploitation started as early as the latest Silurian.

On the enigma of Palaenigma wrangeli (Schmidt), a conulariid with a partly non-mineralized skeleton.

Palaenigma wrangeli (Schmidt) is a finger-sized fossil with a tetraradiate conical skeleton; it occurs as a rare component in fossiliferous Upper Ordovician strata of the eastern Baltic Basin and is known exclusively from north Estonia. The systematic affinities and palaeoecology of P. wrangeli remained questionable. Here, the available specimens of P. wrangeli have been reexamined using scanning electron microscopy and x-ray computed tomography (microCT). Additionally, the elemental composition of the skeletal elements has been checked using energy dispersive X-ray spectroscopy. The resulting 2D-, and 3D-scans reveal that P. wrangeli consists of an alternation of distinct calcium phosphate (apatite) lamellae and originally organic-rich inter-layers. The lamellae form four semicircular marginal pillars, which are connected by irregularly spaced transverse diaphragms. Marginally, the diaphragms and pillar lamellae are not connected to each other and thus do not form a closed periderm structure. A non-mineralized or poorly mineralized external periderm existed originally in P. wrangeli but is only rarely and fragmentary preserved. P. wrangeli often co-occurs with conulariids in fossil-rich limestone with mudstone-wackestone lithologies. Based on the new data, P. wrangeli can be best interpreted as a poorly mineralized conulariinid from an original soft bottom habitat. Here the new conulariinid family Palaenigmaidae fam. nov. is proposed as the monotypic taxon for P. wrangeli.

Fan worms (Annelida: Sabellidae) from Indonesia collected by the Snellius II Expedition (1984) with descriptions of three new species and tube microstructure.

The Indonesian archipelago is one of the most diverse regions in the marine World. Many contributions on polychaete worms have been published since the Dutch Siboga Expedition to the Indonesian archipelago at the end of the 19th century. In this study, we examined specimens of Sabellidae Latreille, 1825 collected during the Snellius II Expedition (1984) to Indonesia, carried out by the Dutch Research Vessel (RV) "Tyro" and the Indonesian RV "Samudera". The results include reports of Acromegalomma acrophthalmos, A. interruptum, A. sp., Bispira manicata, B. porifera, B. secusoluta, Branchiomma boholense, Notaulax pyrrohogaster, N. tenuitorques, N. sp. 3, Parasabella crassichaetae, Perkinsiana anodina, and Sabellastarte spectabilis. In addition, three new species are described: Acromegalomma sumbense sp. nov., Claviramus olivager sp. nov., and Notaulax montiporicola sp. nov., the latter in living coral (Montipora nodosa). Further, Sabella (Potamilla) polyophthalmos Grube is transferred to Pseudopotamilla. Additional histological accounts of B. porifera and tube microstructure of A. acrophthalmos, B. porifera, P. anodina, Pseudopotamilla polyophthalmos and Sabellastarte spectabilis are also included.

Bioerosion of Inorganic Hard Substrates in the Ordovician of Estonia (Baltica).

The earliest bioeroded inorganic hard substrates in the Ordovician of Estonia appear in the Dapingian. Hardgrounds are also known from the Sandbian and Katian. Most of the bioerosion of inorganic hard substrates occurs as the boring Trypanites Mägdefrau, 1932 along with some possible Gastrochaenolites borings. North American hardground borings are more diverse than those in Baltica. In contrast to a worldwide trend of increasing boring intensity, the Estonian record seems to show no increase in boring intensities during the Middle and Late Ordovician. Hardgrounds seem to be more common during the temperate climate interval of the Ordovician calcite sea in Estonia (seven hardgrounds during 15 my) than in the part with a tropical climate (four hardgrounds during 12 my). Bioerosion is mostly associated with carbonate hardgrounds, but cobbles and pebbles broken from the hardgrounds are also often penetrated by Trypanites borings. The general diversity of boring ichnotaxa in Baltica increased from one ichnospecies in the Cambrian to seven by the end of Ordovician, showing the effect of the GOBE on bioeroding ichnotaxa. The diversity of inorganic hard substrate borers increased by only two times. This difference can be explained by the wider environmental distribution of organic as compared to inorganic substrates in the Ordovician seas of Baltica, and their more continuous temporal availability, which may have caused increased specialization of several borers. The inorganic substrates may have been bioreroded only by the generalists among boring organisms.

Evidence of compositional and ultrastructural shifts during the development of calcareous tubes in the biofouling tubeworm, Hydroides elegans.

The serpulid tubeworm, Hydroides elegans, is an ecologically and economically important species whose biology has been fairly well studied, especially in the context of larval development and settlement on man-made objects (biofouling). Nevertheless, ontogenetic changes associated with calcareous tube composition and structures have not yet been studied. Here, the ultrastructure and composition of the calcareous tubes built by H. elegans was examined in the three early calcifying juvenile stages and in the adult using XRD, FTIR, ICP-OES, SEM and Raman spectroscopy. Ontogenetic shifts in carbonate mineralogy were observed, for example, juvenile tubes contained more amorphous calcium carbonate and were predominantly aragonitic whereas adult tubes were bimineralic with considerably more calcite. The mineral composition gradually shifted during the tube development as shown by a decrease in Sr/Ca and an increase of Mg/Ca ratios with the tubeworm's age. The inner tube layer contained calcite, whereas the outer layer contained aragonite. Similarly, the tube complexity in terms of ultrastructure was associated with development. The sequential appearance of unoriented ultrastructures followed by oriented ultrastructures may reflect the evolutionary history of serpulid tube biominerals. As aragonitic structures are more susceptible to dissolution under ocean acidification (OA) conditions but are more difficult to be removed by anti-fouling treatments, the early developmental stages of the tubeworms may be vulnerable to OA but act as the important target for biofouling control.

The earliest giant Osprioneides borings from the Sandbian (late ordovician) of Estonia.

The earliest Osprioneides kampto borings were found in bryozoan colonies of Sandbian age from northern Estonia (Baltica). The Ordovician was a time of great increase in the quantities of hard substrate removed by single trace makers. Increased predation pressure was most likely the driving force behind the infaunalization of larger invertebrates such as the Osprioneides trace makers in the Ordovician. It is possible that the Osprioneides borer originated in Baltica or in other paleocontinents outside of North America.

Endobiotic rugosan symbionts in stromatoporoids from the Sheinwoodian (Silurian) of Baltica.

A paleoecological study of stromatoporoid endobionts was carried out to discern the relationships between symbiotic rugosans and their stromatoporoid hosts. The earliest endobiotic rugosan symbiont Palaeophyllum sp. in Baltica has only been found in the stromatoporoid Ecclimadictyon astrolaxum from Saaremaa, Estonia. The rugosans are vertically oriented inside the stromatoporoid skeleton. Numerous rugosans have their corallites open at the upper, external surface of stromatoporoids, but many are completely embedded within the stromatoporoids. Stromatoporoid hosts were presumably beneficial for rugosans as elevated substrates on a sea floor that offered a higher tier for feeding. Relative substrate stability in the hydrodynamically active shallow waters may have also been beneficial for the rugosans.

Occurrence, formation and function of organic sheets in the mineral tube structures of Serpulidae (polychaeta, Annelida).

A scanning electron microscopy study of organic sheets in serpulid tube mineral structures was carried out to discern their function, formation and evolution. The organic sheets may have some taxonomic value in distinguishing the two major clades of serpulids previously identified. The organic sheets in the mineral tube structure occur only in certain taxa belonging to clade A, but not all species in clade A have them. Organic sheets are best developed in genus Spirobranchus. One could speculate that organic sheets have evolved as an adaption to further strengthen the mechanical properties of the tubes in clade A, which contains serpulids with the most advanced mineral tube microstructures. The organic sheets are presumably secreted with at least some mineral phase.

SEM study of semi-oriented tube microstructures of Serpulidae (Polychaeta, Annelida): implications for the evolution of complex oriented microstructures.

Semi-oriented microstructures represent an intermediate state between isotropic and complex oriented microstructures of serpulids. Semi-oriented microstructures are apparently unique to serpulids, indicating that there may have been unique aspects in the evolution of their biomineralization system. The occurrence of semi-oriented microstructures in serpulids supports the hypothesis that complex oriented microstructures are derived from isotropic microstructures via evolution of a biochemical crystal orientation control mechanism.

On the unique isotropic aragonitic tube microstructure of some serpulids (Polychaeta, Annelida).

A Scanning electron microscopy study of peculiar serpulid tube microstructures was carried out to determine whether this structure is unique to serpulids and to understand its formation and evolution. The isotropic aragonitic tube microstructure of some serpulids, termed spherulitic irregularly oriented prismatic structure (SIOP), is unknown in other animal skeletons. After nucleation, initial spherulitic prismatic aragonite crystals presumably grow in length and width at one end, collide with the surrounding growing crystal fans, and then continue intertwined growth until the secreted carbonate-charged mucus is completely crystallized. Spherulitic irregularly oriented prismati structure presumably evolved either from the IOP or homogeneous rounded crystal microstructures.

Raman spectroscopic study of the mineral composition of cirratulid tubes (Annelida, Polychaeta).

Raman spectroscopy was used to determine the mineralogical composition of the calcareous tubes of three species belonging to the family Cirratulidae. In all three cases, the tubes were found to be aragonitic, confirming previous inferences based on EDX and thin section studies, and corroborated by new EDX analyses revealing the presence of Sr but no Mg. Biomineralization in cirratulids is first recorded in the Oligocene epoch, at a time of aragonite seas. Similarly, the mineralogies of the earliest skeletons matched seawater chemistry in three other polychaete groups that independently evolved calcareous skeletons.