Ecological aspects of a silicified bivalve fauna from the Silurian of Gotland

The silicified Wenlockian (Silurian) bivalve fauna from MÖllbos, Gotland, is part of a life assemblage. The vast number of shells show unusual phenomena, e.g. shell repair, pearl and tumour formation, etc. A number of shells contain epibionts and bored, round holes. Presumptive predators of the bivalve community are discussed. Size-frequency distribution of the two most abundant species possibly reflects age classes. The fauna, comprising eleven species, is dominated by deposit-feeders (90 %). They exhibit niche diversification, including at least three different feeding levels within the sediment.     

Dental wear and oral pathology: possible evidence and consequences of habitual use of teeth in a Swedish Neolithic sample

Atypical wear and oral lesions were studied in the dental remains from the Middle Neolithic Pitted Ware Culture site Ajvide on the island of Gotland in the Baltic Sea. Teeth (n = 764) from 17 males and 11 females were examined microscopically to document unusual wear, assess wear patterns and oral lesions as evidence of habitual behavior, and determine their effect on the oral health of this population. Five atypical wear types were observed: occlusal facets, occlusal excessive load, labial vertical striae, labial horizontal striae, and interproximal striae. Three oral lesions were recorded: chipping, periapical lesions, and dental tilting. Results indicate that teeth were used in a habitual manner at Ajvide, based on the morphology and regularity of the patterns of atypical wear. Differences were observed between the sexes, indicating gender-related differences in the habitual use of teeth. Some wear categories showed a significant correlation with age, signifying increased or accumulated wear with age. Statistically significant positive correlations were found in the molars between occlusal excessive load wear and periapical lesions as well as tilting. Other apparent links were also observed between chipping and vertical striae as well as excessive load, although these were not significant. This suggests a relationship between dental wear and dental pathologies at this site, suggesting that habitual use of teeth indirectly affected the general oral health at the site. Wear patterns, furthermore, seem to mirror both frequent activities as well as single events.     

Silurian Leptaeninae (Brachiopoda) from Gotland, Sweden

Strophomenid brachiopods belonging to the generaLeptaena andLepidoleptaena are described from the uppermost Llandovery — Ludlow succession of Gotland, Sweden. In Gotland,Lepidoleptaena comprises the single speciesL. poulseni, andLeptaena includes four species:L. rhomboidalis, L. sperion, L. depressa andL. parvirugata n. sp.L. depressa shows a considerable amount of morphological variation, which is recognised in the two new subspeciesLeptaena depressa visbyensis n. ssp. from the Llandoverian — Wenlockian Visby Formation, andL. depressa lata n. ssp. from the mid-Wenlockian Slite Group. The distribution of the different species is largely substrate-dependent, withLeptaena rhomboidalis andLepidoleptaena poulseni adapted to high-energy environments with firm substrates, whereas the remaining species preferred fine-grained substrates in low-energy environments.      

Silurian carbonate platforms and extinction events—ecosystem changes exemplified from Gotland, Sweden

Recent and ancient carbonate platforms are major marine ecosystems, built by various carbonate-secreting organisms with different sensitivity for environmental change. For this reason, carbonate platforms are excellent sensors for changes in contemporaneous marine environments. A variety of ecosystem changes in carbonate platforms have previously been recognised in the aftermath of mass extinction events. This paper addresses how two Silurian extinction events among graptolites, conodonts, and pentamerid brachiopods can be related to changes in the style of carbonate production and general evolution of low latitude carbonate platforms in a similar way as previously reported from the major five mass extinctions of the Phanerozoic. Strata formed on Gotland during the Mulde and Lau events share remarkably many similarities but are strikingly different in composition compared to other strata on the island. The event-related strata is characterised by the sudden appearance of widespread oolites, deviating reef composition, flat-pebble conglomerates, abundant micro- and macro-oncoids, stromatolites, and other microbial facies suggesting decreased bioturbation levels in contemporaneous shelf seas. Importantly, these changes can be tied to high-resolution biostratigraphic frameworks and global stable isotope excursions. The anomalous intervals may therefore be searched for elsewhere in order to test their regional or global significance.     

Silurian calcispheres (Calcitarcha) of Gotland (Sweden): Comparisons with calcareous dinoflagellates

Scanning electron microscope examinations of polished and etched surfaces of sediments from the Silurian carbonate platform of Gotland, Sweden, revealed the presence of numerous, morphologically diverse “calcispheres” (Calcitarcha). Some of these spherical calcareous microfossils display wall structures that are surprisingly similar to those of calcareous dinoflagellate cysts. In analogy to the interpretation of the biological affinities of Palaeozoic acritarchs as cysts of organisms that might have been the ancestors of organic-walled dinoflagellates, the Calcitarcha from Gotland can be compared and may possibly be related to organisms that may have been the ancestors of calcareous cyst-producing dinoflagellates that so far have not been observed before the Late Jurassic.     

Large dwellers of the Silurian Halysites biostrome: rhizosessile life strategies of cystiphyllid rugose corals from the Llandovery of Gotland

An exceptionally well‐preserved, unusual biostrome composed of the framebuilding cateniform tabulate coral Halysites catenularius (Linnaeus, 1767) bears an assemblage of the relatively large solitary cystiphyllid rugosan Cystiphyllum visbyense Wedekind, 1927. The corallites of solitary cystiphyllids are embedded within the ranks of the halysitid colonies, which developed on a soft, muddy substrate and in relatively turbid water. The cystiphyllid larvae successively settled mostly on the ranks of halysitid colonies and on colonies of the tiny phaceloid rugose coral Nanophyllum ramosum Johannessen, 1995, whereas calice‐in‐calice recruitment was not successful for these cystiphyllid corals. Further growth of C. visbyense was supported by rhizoid structures, which were most frequently developed on the cardinal (convex) side of the corallite. The process of formation of the rhizoid structures is here discussed and explained in detail, showing that they were formed by the extension of the basal ectodermal tissue of the polyp. The cystiphyllids, which settled on the walls of living corallites of halysitid colonies, used sweeper tentacles to kill the smaller polyps of the colony to maintain the space around them and expand. Hence, they ultimately used the halysitid colonies only as a hard substrate to stabilize their position on the soft muddy sediment.     

Development of facies and C/O-isotopes in transects through the Ludlow of Gotland: Evidence for global and local influences on a shallow-marine environment

Summary The Silurian of Gotland is characterized by repeated changes in depositional facies development. The deposition of uniform sequences of micritic limestones and marls was interrupted four times by the growth of reef complexes and the formation of expanded carbonate platforms. Coinciding with these, often abrupt, facies changes extinction events occurred which predominantly affected nektonic and planktonic organisms. Ratios of carbon- and oxygen-isotopes covary with the facies development. Periods in which the deposition of limestonemarl alternations prevailed are characterized by relatively low C- and O-isotope values. During periods of enhanced reef growth isotope values are high. For these changes,Bickert et al. (1997) assume climatic changes between humid “H-periods”, with estuarine circulation systems and cutrophic surface waters with decreased salinity in marginal seas, and arid “A-periods”, with an antiestuarine circulation and oligotrophic, stronger saline surface waters. In order to separate local and regional influences on the isotopic development from the global trend, the interactions between facies formation and isotope record have to be clarified. For this purpose, the patterns of isotope values in the upper part of the Silurian sequence on Gotland (upper Wenlock —upper Ludlow) has been determined and stratigraphically correlated along four transects through different facies areas. Facies formation during this time interval was investigated by differentiation and mapping of twelve facies complexes in the southern part of Gotland. These include shelf areas, reef complexes with patch reefs and biostromes, backreef facies, and marginal-marine deposits. The good correspondence between the carbon-isotope records of the four transects suggests that local environmental conditions in the different facies areas did not influence the δ¹³C values. Therefore, a supra-regional or even global mechanism for the C-isotope variations is likely. In contrast to carbon istopes, the oxygen-isotope values of the four transects generally show parallel trends, but higher variabilities and in parts distinctly deviating developments with a trend to more negative values. These are interpreted as an effect of local warming in small shallow-water areas which developed during arid periods in reef complexes and backreef areas. The boundaries between A-periods and H-periods, as defined by δ¹³C values, which are interpreted as isochrones, can be mapped. From the upper Homerian to the Pridolian six parastratigraphic isotope zones are defined which only partly match the stratigraphic division ofHede (1942, 1960). The isotope stratigraphy results in an improved correlation between the shallow and marginal-marine areas in the eastern part of Gotland and the uniform shelf areas at the west coast of the island. Furthermore, a detailed relationship between the development of carbon and oxygen isotope ratios, the carbonate facies formation, and the succession of palaeontological events could be observed. At the transition from H-periods to A-periods, major extinction events occurred prior to the first increase of δ¹³C and δ¹⁸O values. Extinction events affected conodonts, graptolites, acritarchs, chitinozoans, and vertebrates and resulted in impoverished nektonic and planktonic communities. The reef-building benthos was less affected. Parallel to a first slight increase of isotope values, facies began to change, and reefs developed in suitable locations. The subsequent rapid increase of C- and O-isotope values occurred contemporarily with strong facies changes and a short-term drop of sea-level. Oligotrophic conditions in the later stages of A-periods led to strong reef growth and to an expansion of carbonate platforms. The transitions from A-periods to H-periods were more gradual. The δ¹³C values decreased slowly, but reef growth continued. Later the reefs retreated and were covered by the prograding depositional facies of shelf areas. The diversity of planktonic and nektonic communities increased again. The close relationship between facies formation, palaeontological events and isotope records in the Silurian suggests common steering mechanisms but gives no indication of the causes for the repeated extincion events related to H-period/A-period transitions. Especially the observation, that strong extinctions occurred prior to changes of isotope values and facies, points to causes that left no signals in the geological record. Hypothetical causes like collapse of trophical nets, anoxias, or cooling events are not evident in the sediment record or do not fit into the regular succession of period transitions.     

Acritarch distribution along an inshore–offshore transect in the Gorstian (lower Ludlow) of Gotland, Sweden

To better understand palaeoecological signatures in Palaeozoic acritarch assemblages, the distribution of palynomorphs has been quantitatively studied in eight localities from the Gorstian, lower Ludlow (Late Silurian) of Gotland, Sweden. The localities are situated along an inshore–offshore transect comprising shallow marine lagoonal environments to distal shelf facies. Process-bearing acritarchs and sphaeromorphs constitute the main components within the palynomorph assemblages. The lateral distribution of palynomorphs exhibits characteristic features at three different levels as follows. (1) With regard to the overall composition of the palynomorph assemblages, the abundance of process-bearing acritarchs increases towards the distal shelf, while the abundance of sphaeromorphs decreases. (2) At the generic level, the acritarchs Micrhystridium and Dilatisphaera are more abundant in the proximal facies, while Evittia, Percultisphaera and Oppilatala are more common in distal environments. (3) At an infrageneric level, Micrhystridium morphotypes with shorter processes are mainly present in proximal environments, while those with longer and ramified processes occur in more distal shelf environments. The palynomorph distribution along the inshore–offshore transect highlights the potential of acritarchs and prasinophytes as palaeoenvironmental indicators.     

Endolithic micro-organisms silicification of a bivalve fauna from the Silurian of Gotland

The silicified Wenlockian bivalve shells at Möllbos have been fragmented to a considerable extent. Shells which were broken prior to silicification exhibit possible original shell layers while those which were fragmented during laboratory treatment show no primary structures. The fauna at Möllbos was attacked by endolithic micro-organisms. The borings of these were then coated with a carbonate envelope. After burial the unattacked original shell material was dissolved the envelope silicified. Later, empty moulds were subsequently filled with drusy calcite occasionally with quartz crystals. A third silicification went occurred at a later diagenetic stage when matrix had become lithified.     

Tracing prehistoric activities: musculoskeletal stress marker analysis of a Stone-Age population on the island of Gotland in the Baltic sea

The skeletal remains from the Middle Neolithic (2750-2300 BC) burial ground at Ajvide, Gotland, are analyzed in order to explore musculoskeletal patterns and to attempt to trace general as well as three specific prehistoric activities (archery, harpooning, and kayaking) that are likely to have been performed in this marine setting of fishing, hunting, and gathering. Scoring of muscular and ligament attachments is performed using the scoring method of Hawkey and Merbs ([1995] Int. J. Osteoarchaeol. 5:324-338) for muskuloskeletal stress markers (MSM). The skeletal material consists of 24 male and 15 female adult individuals divided into three age groups: young (<24 years), middle (25-39 years), and old (>40 years). Thirty upper body MSM sites, on both the left and right sides, are scored and form the basis of the study. Results show that males most frequently have higher mean MSM scores than females. Bilateral asymmetry was noted as low in both sexes. Age proved to be a contributing factor to increased MSM scores, with a greater age-related increase in females. MSM patterns were analyzed statistically in muscle groups associated with the three investigated activities. Significant positive correlations were observed in male individuals in muscle groups associated with archery and to some extent harpooning, an indication that these activities would mainly have been performed by men. Correlations in kayaking muscles were not evidently consistent with the kayaking motion. Furthermore, the costoclavicular ligament, often referred to in connection with "kayaker's clavicle," showed no positive statistical correlation with the kayaking muscles.