The sculpture and histology of osteostracan head shields and trunk squamations from the Silurian of Estonia are described, illustrated and used for the identification of disarticulated microremains collected from outcrop sections and numerous drill cores in Estonia and Latvia over the last 40 years. The scattered osteostracan material contains thousands of specimens of scales and shield fragments. The sculpture and histology of species of the previously identified eight genera in the region (Tremataspis, Dartmuthia, Saaremaaspis, Oeselaspis, Aestiaspis, Thyestes, Procephalaspis, Witaaspis) are redescribed, together with Ateleaspis, found in Estonia for the first time. The sculpture on the cornua of several taxa is described for the first time. The new taxa Tremataspis perforata sp. nov., Dartmuthia procera sp. nov., Eldaaspis miklii gen. et sp. nov., Tahulaspis ordinata gen. et sp. nov., Tahulaspis praevia gen. et sp. nov., Meelaidaspis gennadii gen. et sp. nov. and Ohesaareaspis ponticulata gen. et sp. nov. are established, based upon sculpture and histological characteristics of the exoskeleton. The supraoral field with denticles of that field are described for the first time in Oeselaspis pustulata, as is the supraoral plate with buccal denticles in Meelaidaspis gennadii gen. et sp. nov. Thin sections of all taxa (except of Witaaspis) have been studied. This work sheds light on East Baltic osteostracan biodiversity from the Maasi Beds of the Jaagarahu Stage, Sheinwoodian, lower Wenlock up to the Ohesaare Stage, upper Přidoli, Silurian.

The sculpture of scales and plates of articulated anaspids from the order Birkeniida is described and used to clarify the position of scale taxa previously left in open nomenclature. The dermal skeleton of a well-preserved squamation of Birkenia elegans Traquair, 1898 from the Silurian of Scotland shows a characteristic finely tuberculated sculpture over the whole body. Rhyncholepis parvula Kiær, 1911, Pterygolepis nitida (Kiær, 1911) and Pharyngolepis oblonga Kiær, 1911, from the Silurian of Norway show three other sculpture types. Northern Hemisphere disarticulated scales and plates are described here, supporting a new anaspid taxonomy that includes both articulated and disarticulated material. The diversity, distribution, evolutionary trends and biostratigraphy of anaspids are described in the context of this new taxonomy, which includes six families (two are new) subdivided into 16 genera (10 are new) and 22 species (15 are new).New taxa among Birkeniidae Traquair, 1898 are Birkenia robusta sp. nov. and Hoburgilepis papillata gen. et sp. nov.. Rhyncholepididae Kiær, 1924 includes Rhyncholepis butriangula sp. nov., Silmalepis erinacea gen. et sp. nov., Vesikulepis funiforma gen. et sp. nov., Maurylepis lacrimans gen. et sp. nov., and the previously described Schidiosteus mustelensis Pander, 1856 and Rytidolepis quenstedtii Pander, 1856. Tahulalepididae fam. nov. is represented by Tahulalepis elongituberculata gen. et sp. nov. and the revised T. kingi (Woodward, 1947). Septentrioniidae fam. nov. contains Septentrionia lancifera gen. et sp. nov., S. mucronata gen. et sp. nov., S. dissimilis gen. et sp. nov., S. seducta gen. et sp. nov., Liivilepis curvata gen. et sp. nov., Spokoinolepis alternans gen. et sp. nov. and Manbrookia asperella gen. et sp. nov. The family level position of Ruhnulepis longicostata gen. et sp. nov. is uncertain. Pterygolepididae Obruchev, 1964 and Pharyngolepididae Kiær, 1924 remain monogeneric.

The first vertebrate-based subdivisions of Silurian strata were mainly drawn on material from outcrops in Britain and drill cores from the southern Baltic. Nearly twenty years ago the first vertebrate biozonal scheme was developed on the basis of vertebrate distribution in several continuous drill core sections in the northern Baltic. This paper presents a new scheme in which many new data on vertebrate distribution from the Baltica (Baltic region, NW Russia), Avalonia (southern Britain, eastern Canada), Laurentia (northern Canada, Greenland, Scotland) and Kara (Arctic Russia) palaeocontinents have been used. All the zones have been defined, and the geographical distribution and the reference stratum and locality for each zone have been given. The Llandovery part of the succession contains the Valyalepis crista, Loganellia aldridgei and L. scotica zones; the Wenlock part is represented by the Archipelepis bifurcata/Arch. turbinata, L. grossi, Overia adraini, L. einari and Paralogania martinssoni zones. The Par. martinssoni Zone continues in the Ludlow and is followed by the Phlebolepis ornata, Phl. elegans, Andreolepis hedei, Thelodus sculptilis and T. admirabilis zones. The last zone continues in the lower Přidoli and is followed by the Nostolepis gracilis, Poracanthodes punctatus and Trimerolepis timanica zones. The L. aldridgei and Arch. bifurcata zones are new, and the Arch. turbinata and O. adraini faunas have been raised from ‘faunal succession units’ to zones. The geographically widely distributed L. grossi Zone in the upper Sheinwoodian and the Par. martinssoni Zone in the upper Homerian–lowermost Gorstian allow the integration of regional successions into one Generalized Vertebrate Zonal Scheme. Possible correlations of conodont and vertebrate biozones, and gaps in sedimentation in the northern East Baltic Silurian sequence have been discussed, the most extensive hiatus being between the Paadla and Kuressaare stages

Thyestiids are a group of osteostracans (sister-group to jawed vertebrates) ranging in time from the early Silurian to Middle Devonian. Tremataspis is unique among thyestiids in having a continuous mesodentine and enameloid cover on its dermal elements, and an embedded pore-canal system divided into lower and upper parts by a perforated septum. The origin of this upper mesh canal system and its potential homology to similar canal systems of other osteostracans has remained a matter of debate. To investigate this, we use synchrotron radiation microtomography data of four species of Tremataspis and three other thyestiid genera. Procephalaspis oeselensis lacks an upper mesh canal system entirely, but Aestiaspis viitaensis has partially enclosed upper canals formed between slightly modified tubercles that generally only cover separate pore fields. Further modification of tubercles in Dartmuthia gemmifera forms a more extensive, semi-enclosed upper mesh canal system that overlies an extensive perforated septum, similar to that found in Tremataspis. Lower mesh canals in P. oeselensis are radially arranged and buried tubercles indicate a continuous growth and addition of dermal hard tissues. These features are lacking to varying degrees in the other investigated thyestiids, and Tremataspis probably had a determinate growth accompanied by a single mineralization phase of its dermal hard tissues.The previously proposed homology between the semi-enclosed upper canal system in Dartmuthia to the pore-canal system in Tremataspis is supported in this study, but the suggested homologies between these canals and other parts of the thyestiid vasculature to those in non-thyestiid osteostracans remain unclear. This study shows that three-dimensional modeling of high-resolution data can provide histological and structural details that can help clarify homology issues and elucidate the evolution of dermal hard tissues in osteostracans. In extension, this can give insights into how these tissues relate to those found among jawed vertebrates.

ABSTRACT. Thelodont scales from the Middle Llandovery±lowermost Lochkovian of southern Britain are described with three new taxa, Loganellia? unispinata sp. nov., Nethertonodus prodigialis gen. et sp. nov. and Paralogania tarranti sp. nov. established, and Paralogania kaarmisensis Ma Èrss identi®ed for the ®rst time from Britain. Thelodonts are rare in the Llandovery and Wenlock series where predominantly Loganellia cf. aldridgei Turner, Loganellia scotica (Traquair) and Thelodus sp. are accompanied by coniform conodonts Panderodus Ethington, Decoriconus Cooper and Dapsilodus obliquicostatus (Branson and Mehl). The majority of the material has been recovered from the Ludlow and Pr Ïõ Âdolõ Â series. Thelodus laevis (Pander) and Paralogania martinssoni (Gross) occur in the lower Gorstian. Paralogania kaarmisensis and Phlebolepis elegans Pander come from the Upper Gorstian±Lower Ludfordian and are associated with the zonal conodont Polygnathoides siluricus Branson and Mehl in the upper Gorstian. Thelodus parvidens Agassiz and T. trilobatus Hoppe dominate in the Upper Ludfordian in association with the rarer zonal conodonts Ozarkodina snajdri (Walliser), O. crispa (Walliser) and O. remscheidensis eosteinhornensis (Walliser). The basal Pr Ïõ Âdolõ Â Series indicates a change to a thelodont fauna dominated by Paralogania ludlowiensis (Gross) and including Nethertonodus prodigialis. Higher in the Pr Ïõ Âdolõ Â Series, a succession in faunas includes Katoporodus cf. timanicus (Karataju Åte-Talimaa), Paralogania tarranti, Loganellia? unispinata and Goniporus alatus (Gross). In the highest Pr Ïõ Âdolõ Â beds Paralogania kummerowi (Gross) and Loganellia cuneata (Gross) can be traced before the incoming of the lowermost Devonian taxon Turinia pagei (Powrie) along with Nikolivia gutta Karataju Åte-Talimaa. Correlations with the Baltic are suggested from the Lower Ludlow, Upper Silurian, up to the Lower Lochkovian, Lower Devonian. Gross (1967) studied and described scales from the Ludlow and Temeside Bone Beds and the Psammosteus Limestone, and also scales from some other Devonian outcrops he obtained from O. Walliser of Go Èttingen and T. érvig (1917±1994). Gross (1967) also suggested that British thelodonts were useful for biostratigraphy and correlations with the Baltic and Oslo Region. Turner (1973) gave the distribution of thelodonts from a large number of localities in Britain, recognized a succession of assemblages, and suggested correlations with the Baltic, Scandinavia, Podolia, North Timan, Spitsbergen and Canada. As a result of this study, the distribution of thelodonts was considered when correlating Upper Silurian boundaries in Britain and the Siluro-Devonian boundary in particular (Turner 1977;White and Coppack 1977; Bassett et al. 1982). Thelodonts have been identi®ed from some British borehole studies and used in biostratigraphy (Turner 1973(Turner , 1984(Turner , 2000White and Coppack 1977). New thelodonts were [Palaeontology, Vol. 47, Part 5, 2004 q The Palaeontological Association described during collabo...

Articulated thelodonts, Loganellia scotica (Traquair), Shielia gen.n. taiti (Stetson), Lanarkia horrida Traquair, L. spinulosa Traquair and Turinia pagei (Powrie) from the Silurian and Devonian of Scotland are re-described. A new species, Lanarkia lanceolata sp.n. from the Wenlock, Lower Silurian, is established. For each species, diagnosis is specified, varieties of scale morphology are given, the branchial area is described and body morphology is detailed. A paired ventral fin in Shielia gen.n. taiti and Lanarkia lanceolata sp.n. has been discovered. A new reconstruction of Loganellia and Shielia, and diagnosis for thelodonts from Scotland are given. The locality list of Scottish agnathans is reviewed and the biostratigraphical distribution discussed.

Conodonts, thelodonts and acanthodians have previously been collected from the Přídolí of Shropshire (Welsh Borderland) from bone beds such as the Ludlow Bone Bed. In general, specimens collected from these types of deposit are very abraded and make taxonomic studies difficult. This contribution describes well preserved thelodont microelements from two samples from the Downton Castle Sandstone Formation (Přídolí, Silurian) of the Much Wenlock area of Shropshire. Head scales, a wide variety of transitional scales, and pore scales of the lateral line sensory canals of the thelodont Paralogania ludlowiensis (Gross) are described for the first time. A new acanthodian Nostolepis linleyensis sp. nov. and elements from the apparatus of the conodont Ozarkodina?hemensis sp. nov. are documented from the same samples. The thelodonts in both samples are typical of upper Ludlow–Přídolí faunas from across the Welsh Borderland, and are also present in smaller numbers in the Ludlow Series of Gotland, Sweden, Saaremaa Island, Estonia, and in Latvia and Lithuania. The conodont O? hemensis sp. nov. is restricted to the Much Wenlock area of Shropshire. The co‐occurrence of well preserved elements of conodonts and thelodonts suggests that they were deposited fairly rapidly with little or no re‐working and that a restricted marine environment prevailed in the early Přídolí in the Much Wenlock area of Shropshire.

The numerous cushion-shaped tooth-bearing plates attributed to the stem group osteichthyan Lophosteus superbus, which are argued here to represent an early form of the osteichthyan inner dental arcade, display a previously unknown and presumably primitive mode of tooth shedding by basal hard tissue resorption. They carry regularly spaced, recumbent, gently recurved teeth arranged in transverse tooth files that diverge towards the lingual margin of the cushion. Three-dimensional reconstruction from propagation phase-contrast synchrotron microtomography (PPC-SRµCT) reveals remnants of the first-generation teeth embedded in the basal plate, a feature never previously observed in any taxon. These teeth were shed by semi-basal resorption with the periphery of their bases retained as dentine rings. The rings are highly overlapped, which evidences tooth shedding prior to adding the next first-generation tooth at the growing edge of the plate. The first generation of teeth is thus diachronous. Successor teeth at the same sites underwent cyclical replacing and shedding through basal resorption, producing stacks of buried resorption surfaces separated by bone of attachment. The number and spatial arrangement of resorption surfaces elucidates that basal resorption of replacement teeth had taken place at the older tooth sites before the addition of the youngest first-generation teeth at the lingual margin. Thus, the replacement tooth buds cannot have been generated by a single permanent dental lamina at the lingual edge of the tooth cushion, but must have arisen either from successional dental laminae associated with the individual predecessor teeth, or directly from the dental epithelium of these teeth. The virtual histological dissection of these Late Silurian microfossils broadens our understanding of the development of the gnathostome dental systems and the acquisition of the osteichthyan-type of tooth replacement.