A new microconchid species from the Silurian of Baltica

The diversity of Silurian microconchids is still poorly understood. Here, a new microconchid tubeworm species, Palaeoconchus wilsoni, is described from the Silurian (Ludlow) encrusting rugose corals from Estonia (Saaremaa Island) and a brachiopod shell from Sweden (Gotland). In Estonia, the microconchids are a dominant constituent of the encrusting assemblages, associated with cornulitids, Anticalyptraea, auloporids, trepostome bryozoans, hederelloids and enigmatic ascodictyids. It is notable that these Silurian encrusting assemblages are clearly dominated by tentaculitoids (microconchids, cornulitids and Anticalyptraea) which very often co-exist on the same coral host. Morphologically similar microconchids and Anticalyptraea may have exploited a more similar ecological niche than the straight-shelled cornulitids. However, the clear predominance of microconchids over Anticalyptraea in the communities may indicate that this genus was a less effective competitor for food than microconchid tubeworms

Première description de rares terriers de Teichichnus à partir de roches carbonatées du Paléozoïque inférieur de l'Estonie

Teichichnus burrows occur in the Sandbian, Katian and Telychian of Estonia associated with carbonate rocks. It is possible that Teichichnus is more common in the Sandbian than in the Lower to Middle Ordovician and in the Silurian. Two ichnospecies, T. rectus and T. patens, have been identified from the Lower Paleozoic of Estonia. This is the first record of T. patens in the Ordovician of Baltica. Teichichnus in the Sandbian, Katian and Telychian of Estonia is restricted to the shallowest tier levels. The rarity of Teichichnus in the carbonate sequences of the Ordovician and Silurian of Estonia reflects little bathymetric variability and an extremely low sedimentation rate in the shallow epicontinental basin.Les terriers de Teichichnus sont présents dans le Sandbien, le Katien et le Telychien d'Estonie, associés à des roches carbonatées. Il est possible que Teichichnus soit plus commun dans le Sandbien que dans l'Ordovicien inférieur et moyen ainsi que dans le Silurien. Deux ichno-espèces, T. rectus et T. patens, ont été identifiées dans le Paléozoïque inférieur d'Estonie. Il s'agit du premier enregistrement de T. patens dans l'Ordovicien de Baltica. Teichichnus dans le Sandbien, le Katien et le Telychien d'Estonie est limité aux niveaux les moins profonds. Sa rareté dans les séquences carbonatées de l'Ordovicien et du Silurien en Estonie reflète une faible variabilité bathymétrique combinée à une vitesse de sédimentation extrêmement faible dans le bassin épicontinental peu profond. Mots-clef

Quelques fonds durcis et encroûtés de l'Ordovicien d'Estonie, bouclier balte (Baltica)

The Ordovician hardground faunas of Estonia are not diverse. They include echinoderm holdfasts (i.e., eocrinoids and crinoids), edrioasteroids, bryozoans (both hemispherical trepostomes and stalked ptilodictyids) and cornulitids. The earliest hardground faunas appeared in the Dapingian (i.e., bryozoans and echinoderms). The Estonian hardground faunas are less diverse than the North American ones. North American hardgrounds seem to be more heavily encrusted than the Estonian ones. These differences may be due to the paleogeographic distances, different climates and different sedimentation environments of the paleocontinentsLa faune des fonds durcis ordoviciens est peu diversifiée. On y observe des structures d'ancrage d'échinodermes (d'éocrinoïdes et de crinoïdes), des édrioasteroïdes, bryozoaires (à la fois des trépostomes hémisphériques et des ptilodictyidés pourvus de tiges) et des cornulitidés. Les pre-miers peuplements de fonds durcis apparaissent au Dapingien : il s'agit alors de bryozoaires et d'échinodermes. La faune des fonds durcis estoniens est moins diversifiée que celle d'Amérique du Nord. Les fonds durcis nord-américains semblent être plus fortement encroûtés que ceux d'Estonie. Ces différences tiennent peut-être aux distances entre ces deux paléocontinents ou à des conditions climatiques ou d'environnements de dépôt distinctes

Rare intercroissance tétracoralliaire-bryozoaire dans l'Ordovicien supérieur d'Estonie

Two relatively large specimens of the rugosan Lambelasma sp. are fully intergrown with the bryozoan Stigmatella massalis colony. The intergrown specimen occurs in the Oandu Regional Stage (lower Katian) of Estonia and constitutes the earliest record of bryozoan-rugosan intergrowth from Baltica. Most likely this symbiotic association was accidental. Rugosans presumably benefitted from the bryozoan, which served as an anchor to stabilize them in hydrodynamically active waters. The lack of malformations and no decrease in the size of bryozoan zooids near the rugosans indicate a lack of negative effect of the rugosans on the bryozoan. Bryozoan-rugosan symbiosis is only known from the Ordovician of Baltica and Laurentia.Deux spécimens relativement grands du tétracoralliaire Lambelasma sp. montrent une intercroissance étroite avec une colonie du bryozoaire Stigmatella massalis. Le matériel a été récolté dans l'Étage Régional Oandu (Katien inférieur) d'Estonie et constitue la plus ancienne mention de intercroissance tétracoralliaire-bryozoaire dans l'Ordovicien de Baltica. Très vraisemblablement cette symbiose est accidentelle. Le tétracoralliaire tire probablement bénéfice du bryozoaire dont il se sert comme point d'ancrage pour se stabiliser dans un milieu marin agité. L'absence de malformations et de diminution de la taille des zoécies du bryozoaire au contact du tétracoralliaire indiquent que ce dernier ne perturbe pas le développement du bryozoaire. La symbiose tétracoralliaire-bryozoaire n'est connue que dans l'Ordovicien de Baltica et de Laurentia

Scale dependent diversity of bryozoan assemblages in the reefs of the Late Ordovician Vasalemma Formation, Estonia

The fieldwork for BK and AP was partly funded by the Academy of Finland project ‘Ecological Engineering as a Biodiversity Driver in Deep Time’ (Decision No. 309422). Deutsche Forschungsgemeinschaft (DFG) is appreciated for financial support of AE (project ER 278/10.1). The work is a contribution to the IGCP program 735 ‘Rocks and the Rise of Ordovician Life’.The reefs of the Vasalemma Formation, late Sandbian, Late Ordovician, of northern Estonia contain an exceptional rich and abundant bryozoan fauna. They are an example of contemporaneous bryozoan-rich reefs known from around the world, representing the peak diversification interval of this group during the Ordovician. The global Ordovician bryozoan diversification was associated with a decrease in provinciality, a pattern known from other skeletal marine metazoans of this period. The diversification is associated with climatic cooling and increasing atmospheric and sea water oxygenation. However, the mechanisms that led to the bryozoan diversification are poorly known. Here we estimate the bryozoan richness (α and γ diversity) and turnover (β diversity) at the level of samples, reefs, and formations in the Vasalemma Formation and in contemporaneous reef limestone occurrences of the Baltoscandian region. The resulting richness and turnover values differ among the three observational levels and hence are scale dependent. A consistent pattern with lowest between-reef turnover and relatively high between-sample turnover could be detected, reflecting high small-scale (within reef) heterogeneities in lithology and original bryozoan habitat. This is consistent with published work, in which evidence has been presented for small-scale substrate heterogeneity as the most important diversification driver of the Ordovician brachiopod diversification in the Baltoscandian region. The fact that reefs and their local substrate are strongly organism moderated environments sheds light on the potentially important ecosystem engineering role of organisms, such as bryozoans, for the Ordovician diversification.Publisher PDFPeer reviewe

Ichnogenus Trypanites in the Ordovician of Estonia (Baltica)

Trypanites is a common boring in Ordovician hardgrounds of Estonia (Baltica). The depth of the sedimentary basin and sedimentation rates controlled the distribution of Trypanites. The trace-makersâ community was diverse and changing over time. Three ichnospecies of Trypanites can be distinguished: T. sozialis, T. weisei and Trypanites isp. All three morphotypes can be recognized in the same hardground. It is impossible to distinguish between the different ichnospecies based only on the size of the boring aperture. The depth of early lithification of the seafloor determines the morphological variability seen in T. sozialis. The occurrence of elongated borings, such as T. weisei and Trypanites isp., is related to tropical environments, and their trace-makers strongly preferred substrates with a homogeneous and dense texture. The texture and available volume of hard substrate controls the ichnodiversity of Trypanites ichnospecies

The earliest Ordovician trace fossils Cruziana and Rusophycus from Baltica

Trace fossils of the ichnogenera Cruziana and Rusophycus are described for the first time from the Lower Ordovician of Baltica. These specimens were found from the upper Tremadocianâlower Floian glauconite sandstone of the Leetse Formation in the Leetse and Uuga cliffs on the Pakri Peninsula, North-West Estonia. The one from the Leetse locality was collected already in the 19th century but was hidden in museum collections. On this piece of rock, together with the Rusophycus, brachiopods Leptembolon lingulaeformis and Thysanotos siluricus occur. They indicate that the sample comes from the strata corresponding to the Thysanotossiluricus Brachiopod Biozone of the Hunneberg Stage. Different lithology of the two newly discovered loose slabs with trace fossils found under the Uuga cliff indicates that, most probably, they originate from different levels of the glauconite sandstone exposed in this section. One of them is heavily pyritized and yields a contact surface with the underlying beige argillite of the Varangu Stage with fragments of the graptolite Kiearograptus supremus and some undescribed acrotretid and other linguloid brachiopods. The second, less strongly lithified slab contains abundant fine debris of thin-shelled unidentifiable linguloid brachiopods and probably comes from a higher level. Earlier studies of conodonts revealed that the Prioniodus elegans Conodont Zone is missing in the Uuga section, thus narrowing down the possible interval of origin of these ichnotaxa to the Paroistodus proteus zone. Interestingly, these two slabs preserve the dissimilar pattern of grouping and orientation of the multiple Rusophycus/Cruziana traces giving some idea about the ethology of trilobites who probably left these traces. The earliest trilobites in the Ordovician succession of Estonia are recorded from the Mäeküla Member, the uppermost part of the Leetse Formation, from an interval where calcareous component first appears in the sediment and thus also the trilobites with their calcitic exoskeleton are preserved. The only trilobites recorded from the Mäeküla Member of the Leetse Formation in these two localities are specimens of Paramegistaspisleuchtenbergi who could have been the trace maker with its macropygidium being of similar size to its cephalon if the second slab would come from the same interval. However, there are more candidates, mainly isoteline trilobites with similar characteristics which are preserved in older but calcareous succession in Sweden and Norway

Répartition de Conichnus et d'Amphorichnus dans le Paléozoïque inférieur d'Estonie (bouclier balte)

Conichnus conicus and Amphorichnus papillatus occur in clay-rich carbonate rocks in the Ordovician of Estonia. Conichnus conicus also occurs in clay-rich carbonates of the early Silurian of Estonia. Lateral adjustment traces are more common in C. conicus than previously recorded. The lack of adjustment traces in Amphorichnus, together with its morphology, does not support synonymy of Conichnus and Amphorichnus. The Conichnus conicus and Amphorichnus papillatus tracemakers preferred shallow water carbonate environments with high clay content. They were rare or did not occur in deeper water muddy environments or where shallow water carbonates accumulated. A high content of volcanic ash in the depositional environment is characteristic of both the Ordovician and Silurian maxima of Conichnus conicus occurrence. C. conicus may have been more common in the temperate seas of Baltica than in the tropics.Conichnus conicus et Amphorichnus papillatus sont présents dans des roches carbonatées riches en argiles de l'Ordovicien d'Estonie. Conichnus conicus est également présent dans des roches carbonatées riches en argiles du Silurien inférieur d'Estonie. Les signes d'ajustements latéraux sont plus fréquents chez C. conicus que ce que nous connaissions auparavant. L'absence de signes d'ajustements chez Amphorichnus, conjointement à sa morphologie, ne vient donc pas corroborer la synonymie de Conichnus et d'Amphorichnus. Les organismes à l'origine des traces de type Conichnus conicus et Amphorichnus papillatus préféraient des environnements peu profonds, carbonatés mais présentant une teneur élevée en argile. Ils sont, par contre, rares ou absents dans des environnements peu profonds mais où les carbonates s'accumulent ou dans des environnements plus profonds et boueux. Une forte teneur en cendres volcaniques dans l'environnement de dépôt représente un trait caractéristique des pics d'abondance ordoviciens et siluriens de Conichnus conicus. C. conicus semble avoir été plus fréquent dans les mers tempérées du bouclier balte que dans celles sous les tropiques

Parasite-induced shell damage in brachiopod Porambonites (Porambonites) laticaudata from the Late Ordovician (Sandbian) of Estonia

A new type of shell damage has been described in Ordovician brachiopods in Porambonites (Porambonites) laticaudata. There is a pair of small pits with somewhat different outline in the shell surface at the anterior commissure of the brachiopod. These pits are oriented in lateral direction, about 40o from the direction of the sulcus on the anterior commissure. Previously known shell damage has resulted from failed predatory attacks by durophagous predators and differ from the shell damage in P. (P.) laticaudata. The pits in the shell margin are most likely the result of shell malformation caused by the presence of symbionts. It is plausible that the symbionts of the P. (P.) laticaudata benefitted from inhalant currents and were cleptoparasites. The symbionts caused damage to the host brachiopod, which also suggests a parasitic relationship

Tremichnus dans des segments de tiges de crinoïdes du Silurien d'Estonie occidentale (bouclier balte)

Rare pits attributed to Tremichnus have been found in crinoids from the Silurian of Estonia. The Rhuddanian Tremichnus is the earliest symbiont in crinoid columnals of Baltica. These pits presumably were domiciles of unknown organisms. Tremichnus had a negative effect on the host crinoid as demonstrated by swollen columnals. Tremichnus in the Silurian of Estonia is less common than similar traces in the Silurian of nearby Gotland. The most important aspect of this study is the rarity of this interaction in these samples in contrast to most other samples of comparable age elsewhere. These structures have a very patchy distribution