The origin of tetraradial symmetry in cnidarians

Serially arranged sets of eight septa‐like structures occur in the basal part of phosphatic tubes of Sphenothallus from the early Ordovician (early Floian) Fenxiang Formation in Hubei Province of China. They are similar in shape, location and number, to cusps in chitinous tubes of extant coronate scyphozoan polyps, which supports the widely accepted cnidarian affinity of this problematic fossil. However, unlike the recent Medusozoa, the tubes of Sphenothallus are flattened at later stages of development, showing biradial symmetry. Moreover, the septa (cusps) in Sphenothallus are obliquely arranged, which introduces a bilateral component to the tube symmetry. This makes Sphenothallus similar to the Early Cambrian Paiutitubulites, having similar septa but with even more apparent bilateral disposition. Biradial symmetry also characterizes the Early Cambrian tubular fossil Hexaconularia, showing a similarity to the conulariids. However, instead of being strictly tetraradial like conulariids, Hexaconularia shows hexaradial symmetry superimposed on the biradial one. A conulariid with a smooth test showing signs of the ‘origami’ plicated closure of the aperture found in the Fenxiang Formation supports the idea that tetraradial symmetry of conulariids resulted from geometrical constrains connected with this kind of closure. Its minute basal attachment surface makes it likely that the holdfasts characterizing Sphenothallus and advanced conulariids are secondary features. This concurs with the lack of any such holdfast in the earliest Cambrian Torellella, as well as in the possibly related Olivooides and Quadrapyrgites. Bilaterally arranged internal structures in polyps representing probably the oldest medusozoans support the suggestions based on developmental evidence that the ancestor of cnidarians also was a bilaterally symmetrical animal. This is one more example of fossil data that strictly fit the molecular phylogenetic evidence but not necessarily morphology‐based zoological interpretations.     

Evolution within the Silurian cephalopod genusInversoceras

Specimens of the Silurian cephalopod genusInversoceras from the Wenlockian Slite Beds of Gotland, Sweden, are more curved and swollen than those from the Ludlowian Hemse Beds of Gotland. The former population is interpreted as a direct ancestor to the latter. The younger form,Inversoceras p. perversum, originally described by Barrande from Bohemia, includes both individuals with a fairly straight shell and with a slightly curved shell. The distinctly curved and swollen form from the Wenlock beds is herein described as a new subspecies,I. p. eoperversum. I. p. falciformis from the Ludlowian of Bohemia is excluded fromI. perversum and. treated as a distinct species. Its relationship to the olderI. p. eoperversum is discussed. This discussion also includesI. dayi from the Niagarian of North America.     

Silurian craniide brachiopods from Gotland

Seven craniide brachiopod genera are described from the Silurian (Wenlock–Ludlow) of Gotland, including one new genus and five new species. The new genus and species Thulecrania septicostata is unique among Silurian craniides as it possesses solid spines. The new species Lepidocrania multilamellosa is the first Silurian record of this poorly known Permian genus. The problematic North American Propatella Grubbs, 1939 , was originally described as a gastropod, but the new species Propatella palmaria from Gotland shows that it is a craniide with sutured hollow spines of a type not previously recorded from Silurian craniides. The dorsal valves of the new species Valdiviathyris? bicornis are remarkably similar to those of the type species and represent the first possible Palaeozoic record of this poorly known extant craniide. This first systematic study of craniide brachiopods from the Silurian of Gotland shows that the diversity is relatively high as compared to other known Silurian craniide faunas, but a more thorough comparison is not possible due to the lack of data from most parts of the world. The new data from Gotland support the view that the craniides were not affected by the end‐Ordovician extinction.     

Patterns of sclerobiont colonization on the rugose coral Schlotheimophyllum patellatum (Schlotheim, 1820) from the Silurian of Gotland,Sweden

Analysis of mushroom-shaped rugose corals Schlotheimophyllum patellatum (Schlotheim, 1820) from the Silurian (Upper Visby Beds, Lower Wenlock, Sheinwoodian) of Gotland, Sweden, showed that they were colonized on both the upper (exposed) and lower (cryptic) sides by a variety of encrusting and boring (sclerobiont) biotas, represented by 10 taxa and at least 23 species. Bryozoans and microconchid tubeworms, the most abundant encrusters, dominated on the cryptic undersides of the corals, while the dominant endobionts responsible for Trypanites borings overwhelmingly dominated the exposed surfaces. Except for cnidarian sphenothallids, which were exclusive colonizers of the underside of only one coral host, no other encrusters could be referred to as obligate cryptobionts. Because the upper surface of these corals was likely covered by soft-tissues during life, in specimens lifted off the sea-floor sclerobionts must have settled on the cryptic sides first. They could colonize the upper side only after the coral’s death, unless it was covered by sediment as could be the case in some flat specimens. With time, the space on the underside of the coral skeleton may have progressively been filled by sediment as well, precluding further colonization by sclerobionts. In that respect, the colonization patterns of these corals by encrusters and borers were controlled by the complex interplay of environmental factors, sclerobiont dynamics and coral growth in a given Silurian habitat. Compared with Silurian stromatoporoid hosts, the sclerobiont diversity and abundance noted on the Schlotheimophyllum corals may be regarded as representative for the Silurian as a whole.     

Earliest ontogeny of Early Palaeozoic Craniiformea: compelling evidence for lecithotrophy

Popov, L.E., Bassett, M.G. & Holmer, L.E. 2012: Earliest ontogeny of Early Palaeozoic Craniiformea: compelling evidence for lecithotrophy. Lethaia, Vol. 45, pp. 566–573. The early ontogeny of Palaeozoic Craniiformea (Brachiopoda) remains controversial, with conflicting reports of evidence indicating lecithotrophic versus planktotrophic larval stages. Further compelling evidence for lecithotrophy in Palaeozoic craniiforms is described here. Newly obtained, well‐preserved Silurian specimens of craniiforms, including Craniops (Craniopsida), and Lepidocrania? and Orthisocrania (Craniida) from Gotland and the St. Petersburg region, form the basis for this study. The new material demonstrates that the characters of shell structure and shell formation provide evidence of early differentiation of an adult dorsal mantle, and the presence of a distinctive primary layer with a characteristic lath‐like pattern indicates that these craniiforms underwent a lecithotrophic larval stage, more or less identical to that of living. □Novocrania. Brachiopoda, Craniiformea, ontogeny, phylogeny, Early Palaeozoic.     

Silurian scolecodonts

Bergman, Claes F. 1989: Silurian Paulinitid Polychaetes from Gotland. Fossils and Strata. No. 25.     

Paleoecology of a low-diversity silurian community from the tofta beds of Gotland

The Silurian (Wenlockian) Tofta Beds at Galgberget 1, Gotland, Sweden, formed in a protected intertidal setting. Massive fenestral limestone at this locality contains a low diversity community dominated by stromatoporoids, calcareous algae, and ostracods, with less common rugose corals, bryozoans, brachiopods, and trilobites. Abundance of stromatoporoids, which form about 40% of sediment volume, suggests reef-like conditions. The Tofta community differs from typical Silurian reef communities, however, in its low diversity, very limited tiering, and absence of groups such as crionozoans and tabulates. These differences are possibly due to intertidal conditions which precluded upward growth of a mound structure and subjected the community to periodic desication.     

Silurian (Ludlow) acastid trilobites from Gotland and Scania,Sweden

Acastid trilobites from Ludlow strata on Gotland include three stratigraphically non-overlapping species ofAcastella Reed:Acastella madidipes n. sp. (Hemse Marl and Eke Beds),A. breviceps (Angelin 1851) (Burgsvik Beds), andA. amatrix n. sp. (Hamra Beds, and probably Burgsvik Beds). Type and other material ofA. breviceps (Angelin) from the Klinta Formation, Scania, is revised. Meraspid and early holaspid material ofAcastella is described. Some characters supportAcaste and allied taxa of mostly Wenlock age as a monophyletic subfamily Acastinae.Acastella as widely recognized is a grade with Devonian species more closely related to Asteropyginae than to the Ludlow type species. Baltic Ludlow-P?00ED;dolí species previously placed inScotiella Delo form a distinct group,Ewacaste n. gen., that includes a species from the uppermost Eke Beds on Gotland.     

LOWER SILURIAN THELODONTS FROM PRINCE OF WALES ISLAND, NORTHWEST TERRITORIES

Thelodont scales, determined as Thelodus laevis (Pander), Logania martinssoni Gross, and Phlebolepida gen. et sp. indet., and other agnathan remains have been found in the dolomite-limestone sequence of the Transition Beds between the Allen Bay and Read Bay Formations. This thelodont association is typical of the Middle and Lower Upper Silurian of Ösel and Gotland, and its presence therefore provides evidence for the upper age limit of the transition beds.     

Cord‐forming Palaeozoic fungi in terrestrial assemblages

The fossil record paints a thin picture of early terrestrial life. Useful diagnostic features are rare in the organic‐walled fossils of the first land colonizers, and at first glance the Silurian–Devonian Tortotubus protuberans seems no exception. Now, new material from New York, Gotland and Scotland reveals the ontogenesis and affinity of this problematic organism. Its filamentous early stages (previously referred to Ornatifilum lornensis) demonstrate simple septal perforations and a bilayered cell wall; threads of entwined filaments, bounded by an elaborately sculptured surface, arose via the retrograde growth and subsequent proliferation of secondary branches. This morphology and pattern of growth together indicate an affinity with the ‘higher’ fungi (Dikarya) and document the formation of differentiated mycelium. The presence of complex mycelial fossils in the earliest Silurian corroborates the likely contribution of fungi to the colonization of land and the establishment of modern sedimentological systems; their rise seemingly accompanied the diversification of early embryophytes and the vegetation of the terrestrial biosphere.     

Revised correlation of Silurian Provincial Series of North America with global and regional chronostratigraphic units and δ13Ccarb chemostratigraphy

Cramer, B.D., Brett, C.E., Melchin, M.J., Männik, P., Kleffner, M.A., McLaughlin, P.I., Loydell, D.K., Munnecke, A., Jeppsson, L., Corradini, C., Brunton, F.R. & Saltzman, M.R. 2011: Revised correlation of Silurian Provincial Series of North America with global and regional chronostratigraphic units and δ¹³C_carb chemostratigraphy. Lethaia, Vol. 44, pp. 185–202. Recent revisions to the biostratigraphic and chronostratigraphic assignment of strata from the type area of the Niagaran Provincial Series (a regional chronostratigraphic unit) have demonstrated the need to revise the chronostratigraphic correlation of the Silurian System of North America. Recently, the working group to restudy the base of the Wenlock Series has developed an extremely high‐resolution global chronostratigraphy for the Telychian and Sheinwoodian stages by integrating graptolite and conodont biostratigraphy with carbonate carbon isotope (δ¹³C_carb) chemostratigraphy. This improved global chronostratigraphy has required such significant chronostratigraphic revisions to the North American succession that much of the Silurian System in North America is currently in a state of flux and needs further refinement. This report serves as an update of the progress on recalibrating the global chronostratigraphic correlation of North American Provincial Series and Stage boundaries in their type area. The revised North American classification is correlated with global series and stages as well as regional classifications used in the United Kingdom, the East Baltic, Australia, China, the Barrandian, and Altaj. Twenty‐four potential stage slices, based primarily on graptolite and conodont zones and correlated to the global series and stages, are illustrated alongside a new composite δ¹³C_carb curve for the Silurian. Conodont, graptolite, isotope, New York, Ontario, series, Silurian, stage.     

Microplankton changes through the early Silurian Ireviken extinction event on Gotland, Sweden

This paper documents and analyses the extinction and origination patterns of acritarchs and prasinophyte algae at the Llandovery/Wenlock boundary transition in the Lower Silurian on the island of Gotland, Sweden. Closely spaced samples were collected from two parallel sections: Lusklint 1 and Lickershamn 2, spanning the upper part of the Lower Visby Beds and almost all of the Upper Visby Beds (i.e. the uppermost Llandovery and lowermost Wenlock).

At least eight extinctions affecting the conodont record have been reported at these levels. This turnover (named the Ireviken Event) has been interpreted as an example of the change from a P to an S climate state, reflecting large changes in the ocean/atmosphere system.

The palynomorph data show a significant turnover in the phytoplankton, with most of the extinctions at the end of the event (85.3% in the top 4 m of the Lusklint 1 section). The originations are more numerous than the extinctions and they are distributed through the whole of the Ireviken Event. There is an uneven distribution across the event with more originations in the Lower Visby Beds forming a convex pattern.

Comparison of these data to other palynological studies suggests that there was a slightly higher number of migrations out of Gotland than in. The P and S model does not successfully explain all the changes recorded, but is the model most inclusive of all the climatic variables available at this time.     

Paleoecologic,temporal, and spatial analysis of early Silurian Reefs of the Chicotte Formation,Anticosti Island,Quebec, Canada

Summary Reefs of the Lower Silurian Chicotte Formation are the largest and most faunally diverse known on Anticosti Island, Quebec. They reach up to 25 m in thickness and 250 m in diameter and are present predominantly at two intervals, forming a lower and upper reef cluster. Remnants of bioherms are represented on the present-day wave-cut terrace as 60 to 100 m diameter, subcircular erosional depressions known as Philip structures or as outcrop. The bioherms were relatively low structures, with approximately 3 to 5 m maximum synoptic relief, some of which developed on hardgrounds and possible paleokarst surfaces of crinoidal wackestone and packstone. Dominant skeletal framework builders and sediment producers within all of the reefs are laminar to low domical stromatoporoids, colonial cerioid and fasciculate rugose corals, colonial tabulate corals, and cryptostome bryozoans. Vertical zonation of reef biota is evident within well-exposed reefs of the lower reef cluster. Three to four stages are recognizable:1) a low-diversity tabulate coral-dominatedpioneering community including large tabulate coral colonies (halysitids and favositids), and few stromatoporoids (clathrodictyids, ecclimadictyids), fasciculate rugosans, large generally monotypic stalked crinoids, and shelly benthos (brachiopods, few ostracodes and trilobites);2) an intermediate- to high-diversity, mixed tabulate coral-stromatoporoid-dominatedreef-core community;3) a slightly lower diversity stromatoporoid-tabulate coral-dominatedclimax community with laminar coenitids and alveolitids; and,4) in a few localities, a capping, low-diversity tabulatecoral-dominated (alveolitid and coenitid), and stromatoporoid-bearing community comprising laminar forms. Amelioration of Early Silurian climates, following Late Ordovician glaciation, allowed gradual reestablishment of extensive shallow-water reef growth, by mainly new and increasingly diverse genera and species of metazoans. Reef development within the Chicotte Formation coincided with global, widespread development of latest Llandovery and earliest Wenlock reefs in subtropical to tropical areas. Chicotte reefs have broad characteristics, in terms of overall biotic composition, vertical successions recognized, and paleogeographic setting, similar to those of equivalent and slightly younger age from intracratonic settings in Baltica (Gotland, Sweden and Estonia) and central and northern Laurentia (Midcontinent, U.S.A.; Hudson Bay, Canada; and North Greenland, Denmark).     

Coralline red algae from the Silurian of Gotland indicate that the order Corallinales (Corallinophycidae,Rhodophyta) is much older than previously thought

Aguirrea fluegelii gen. et sp. nov. (Corallinales, Corallinophycidae, Rhodophyta) is described from the mid‐Silurian of Gotland Island, Sweden (Högklint Formation, lower Wenlock). The holotype is of dimerous construction and includes a uniporate conceptacle with a sporangium, thus providing evidence that taxa of the Corallinales/Corallinaceae existed at least 300 million years earlier than previously documented. Aguirrea fluegelii cannot be unequivocally placed in any of seven currently recognized lineages/subfamilies/groups of the Corallinaceae as not all diagnostic characters are preserved, and thus is accorded incertae sedis status within the family Corallinaceae and order Corallinales. Extant evolutionary history studies of Corallinophycidae involving molecular clocks now require updating using new calibration points to take account of the much earlier unequivocal mid‐Silurian record of uniporate conceptacle‐bearing taxa of Corallinales/Corallinaceae as well as the parallel record of Graticula, a genus attributed to the Sporolithales.     

Astogenese vonAulopora cf.enodis Klaamann 1966Visby-Mergel, Silur von Gotland

This paper deals with bifurcation angle of a reptant, dichotomously branchingAulopora cf.enodis colony (Silurian, Gotland). It can be demonstrated, that during early astogenetic growth the branching angle between offsetting corallites decreases rapidly from 165° to 100° averagely during later growth stages. This growth pattern resembles development of similar dichotomous branching uniseral bryozoans, e.g. PaleozoicCorynotrypa species or MesozoicStomatopora species.     

Earliest ontogeny of the Silurian orthotetide brachiopod Coolinia and its significance for interpreting strophomenate phylogeny

Well‐preserved juvenile specimens of the orthotetide brachiopod Coolinia pecten (Linnaeus, 1758 ) from the Silurian of Gotland, Sweden, demonstrate evidence of a planktotrophic larval habit. Larval shell morphology indicates the absence of a pedicle sheath: this character is otherwise typical of derived billingsellides, strophomenides and productides, which form the conventional strophomenide clade. The presence of a rudimentary colleplax structure in the larval shell of Colinia suggests instead a phylogenetic link to chiliate brachiopods and the enigmatic genus Salanygolina. This relationship suggests an early divergence of rhynchonellate and strophomenate brachiopods.     

Application of Multivariate Analysis for the Selection of Candidates for Biological Control Agents

A survey of natural enemies ofSenna obtusifolia(Leguminosae: Cesalpinaceae) was made in 46 localities in Brazil. Twenty-one preselected insect species were used in a multivariate analysis in a search for potential biocontrol agents, based on frequency of occurrence of each species, degree of insect/host association, and damage level. Insect communities were organized in a binary matrix (presence/absence of the species); measures of distance and similarities were based on the Jaccard coefficient, and cluster analysis was then conducted. Construction of a dendogram showed similarities among geographically close localities where phenological stages of sicklepod plants were similar. Analysis of correspondence based on the correlation matrix showed that Eigen values of the three first axes explain only 40.1% of the observed variation. Correlation among the scores produced by species Eigenvectors in each locality on axis 1 and the latitude, altitude, and phenological stage showed that only phenological stage was correlated (r = 0.45,P = 0.02). The results indicate that weather will not be a limiting factor for the establishment of these insect species in an environmental gradient similar to that of the surveyed area (12 to 22° latitude and 6 to 1200 m altitude). Distribution in the ordinal space of axes 1 and 2 showed thatAgrilus oceanicum(Buprestidae),Fundella argentina(Pyralidae),Thyphedanus undulatus(Hesperidae), andPhoebis sennae(Pieridae) are grouped and separated from the others in axis 2, suggesting that these species co-occur with each other. Based on their feeding guilds, multiple introduction of these species can be considered. Multivariate analysis showed potential to be used as an additional tool to help in the selection of candidates for biocontrol agents.     

The Pleistocene archaeology and environments of the Wasiriya Beds, Rusinga Island, Kenya

Western Kenya is well known for abundant early Miocene hominoid fossils. However, the Wasiriya Beds of Rusinga Island, Kenya, preserve a Pleistocene sedimentary archive with radiocarbon age estimates of >33–45 ka that contains Middle Stone Age artifacts and abundant, well-preserved fossil fauna: a co-occurrence rare in eastern Africa, particularly in the region bounding Lake Victoria. Artifacts and fossils are associated with distal volcanic ash deposits that occur at multiple localities in the Wasiriya Beds, correlated on the basis of geochemical composition as determined by electron probe microanalysis. Sediment lithology and the fossil ungulates suggest a local fluvial system and associated riparian wooded habitat within a predominantly arid grassland setting that differs substantially from the modern environment, where local climate is strongly affected by moisture availability from Lake Victoria. In particular, the presence of oryx (Oryx gazella) and Grevy’s zebra (Equus grevyi) suggest a pre-Last Glacial Maximum expansion of arid grasslands, an environmental reconstruction further supported by the presence of several extinct specialized grazers (Pelorovis antiquus, Megalotragus sp., and a small alcelaphine) that are unknown from Holocene deposits in eastern Africa. The combination of artifacts, a rich fossil fauna, and volcaniclastic sediments makes the Wasiriya Beds a key site for examining the Lake Victoria basin, a biogeographically important area for understanding the diversification and dispersal of Homo sapiens from Africa, whose pre-Last Glacial Maximum history remains poorly understood.     

Some Silurian Brachiopods from Lithuania and Their Palaeobiogeographical Significance

Silurian brachiopods are described from 21 boreholes in Lithuania. Nine genera are recognized and represented by one or two species, including the atrypids Atrypoidea, Cromatrypa, Lissatrypa and Septatrypa, athyridids Collarothyris, Nucleospira, Meristina and Pseudoprotathyris, and rhynchonellids Ancillotoechia and Plagiorhyncha. Two new species, Cromatrypa? pubes and Lissatrypa lithuanica, are described. The same taxa are reported from England, Gotland, Belarus and Podolia in Wenlock and Ludlow strata, and from Podolia, Urals, the Canadian Arctic archipelago and Russian Arctic islands in Prídolí strata.     

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.