Ordovician and Silurian coralline red algae

Solenopora gotlandica Rothpletz, hitherto placed in the Solenoporaceae Pia, is locally common in the mid-Silurian of Gotland and Wales. It possesses small cells, radial monomerous thallus organization, trichocytes, and sporangia arranged in irregular sori. Solenopora gotlandica is not congeneric with S. spongioides , the type species of Solenopora. It closely resembles the extant coralline red alga Sporolithon Heydrich. S. gotlandica is here transferred to Craticula gen. nov., and Craticula is placed in the Craticulaceae fam. nov., which morphologically closely resembles the Mesozoic-Cenozoic family Sporolithaceae. The Craticulaceae is placed in the extant rhodophyte order Corallinales. Middle-Late Ordovician Petrophyton kiaeri Høeg resembles Late Ordovician Solenopora richmondense Blackwell et al. , which has sporangia in calcified compartments arranged in sori. These fossils may be craticulaceans. The apparent gap in the fossil record between Silurian Craticula and Cretaceous Sporolithon favours separation of these genera at family level. However, future work may demonstrate that the Craticulaceae is a junior synonym of the Sporolithaceae. Recognition of the Craticulaceae extends the earliest record of the Corallinales from Cretaceous to Silurian, and possibly Ordovician.     

The role of epistatic interactions underpinning resistance to parasitic Varroa mites in haploid honey bee (Apis mellifera) drones

The Red Queen hypothesis predicts that host–parasite coevolutionary dynamics can select for host resistance through increased genetic diversity, recombination and evolutionary rates. However, in haplodiploid organisms such as the honeybee (Apis mellifera), models suggest the selective pressure is weaker than in diploids. Haplodiploid sex determination, found in A. mellifera, can allow deleterious recessive alleles to persist in the population through the diploid sex with negative effects predominantly expressed in the haploid sex. To overcome these negative effects in haploid genomes, epistatic interactions have been hypothesized to play an important role. Here, we use the interaction between A. mellifera and the parasitic mite Varroa destructor to test epistasis in the expression of resistance, through the inhibition of parasite reproduction, in haploid drones. We find novel loci on three chromosomes which explain over 45% of the resistance phenotype. Two of these loci interact only additively, suggesting their expression is independent of each other, but both loci interact epistatically with the third locus. With drone offspring inheriting only one copy of the queen's chromosomes, the drones will only possess one of two queen alleles throughout the years‐long lifetime of the honeybee colony. Varroa, in comparison, completes its highly inbred reproductive cycle in a matter of weeks, allowing it to rapidly evolve resistance. Faced with the rapidly evolving Varroa, a diversity of pathways and epistatic interactions for the inhibition of Varroa reproduction could therefore provide a selective advantage to the high levels of recombination seen in A. mellifera. This allows for the remixing of phenotypes despite a fixed queen genotype.     

Radial perforations in crinoid stems from the Silurian of Gotland

Radiating intracolumnal canals are a characteristic feature of large (diameter 10 mm or more) crinoid stems from the Silurian of Gotland. They are found in nodals as well as in internodals where the columnal height exceeds one millimetre. They were formed secondarily in the median and distal portions of crinoid stems with pseudocirriferous holdfasts. Intercolumnal canals are found in the distal parts of stems with true cirri regardless of the size of the stem. It is suggested that these canals played an important role in crinoid physiology. The crinoids are believed to have sustained a large proportion of their tissues through cutaneous digestion and uptake of dissolved substances from the surrounding sea water. The intra- and intercolumnal canals increased the surface of the axial canal in relation to volume. They provided a connection between the axial canal and the surrounding sea water, thus facilitating nutrient transport to the tissues.     

Spatial competition among clonal organisms in extant and selected paleozoic reef communities

Summary Occurrences of densely packed benthic organisms in extant reefs are of two types: 1) live-live interactions, where two living organisms interact, and 2) live-dead associations, where only one is alive and uses the other as a substrate. The latter are common in reef deposits due to biostratinomic feedback, i.e. dense skeletal accumulations provide hard substrates for clonal recruitment, thus facilitating greater frequency of live-dead encounters than in lower biomass level-bottom communities dominated by solitary organisms. Differentiating between these two types in ancient reefs is difficult, often impossible. Most live-live interactions among clones in extant reef communities involve competition for space. Clonal spatial competition is divisible into four types: 1) direct-aggressive: encrusting overgrowth; 2) indirect-passive: depriving neighbors of resources, chiefly sunlight, by growth above them; 3) stand-off: avoidance of competition by organisms adopting positions that avoid or minimize direct polyp/zooid contact; and 4) overwhelming: one clone/ species volumetrically or numerically overwhelms the other, meeting minimal resistance. Despite class-order level differences in taxa, our results indicate that extant analogs, based on the arrangement and distortion of skeletons, are valuable for recognizing live-live interactions in Silurian and Carboniferous reefs and interpreting the types of spatial competition represented. Comparison of overhead (plan) views of live-live coral competition in Polynesian reefs with vertical sections of Silurian and Carboniferous sponge-dominated reefs and biostromes suggests that direct-aggressive competition is more common among extant than among Paleozoic reef-builders. Stand-offs showing clone margin distortion and overwhelming with minor skeletal distortion are most common in our fossil examples and probably relate to the dominance of these reefs by sponges. Success by extant sponges in spatial competition is largely due to allelochemical deterrence which may explain the predominance of stand-off and overwhelming confrontations in fossil sponges rather than tentacle-mesentery based direct aggression among extant corals and bryozoans.     

A sequence stratigraphical model for the Late Ludfordian (Silurian) of Gotland,Sweden: implications for timing between changes in sea level,palaeoecology, and the global carbon cycle

This paper investigates a time interval within the Late Ludfordian (Late Silurian), involving changes in faunal composition (the Lau Event), a major positive carbon isotope excursion (CIE), and contemporaneous sea-level changes in remote palaeo-basins. Based on the Silurian strata of Gotland (Sweden), we integrate sequence stratigraphy, carbon isotope stratigraphy, and platform-scale palaeoecological changes associated with this turbulent time period in Earth history. Three depositional sequences (sequences Nos. 1–3), including two separate periods of forced regression (falling stage systems tracts, FSSTs) are identified from outcrop and drillcore studies. The sequence stratigraphical framework is interpreted to reflect glacio-eustatic sea-level changes. The CIE starts at the onset of the initial FSST (sequence No. 1), just below the last appearance datum of the conodont Polygnathoides siluricus. The values increase through the ensuing lowstand and transgressive systems tracts (LST and TST) of sequence No. 2 and peak in the following highstand systems tract (HST). A second forced regression (FSST of sequence No. 2) took place in the lower Ozarkodina snajdri Zone. δ¹³C data are scarce from these siliciclastic strata, but inferably remain high. The δ¹³C values increase within the LST and earliest TST of sequence No. 3, before a decreasing trend starts within the early TST. δ¹³C values return to pre-excursion levels within the ensuing HST. The CIE is closely associated with an increase in stromatolites (mats and oncoids) across a wide range of depths and sedimentary environments, and correlations to other basins indicate a global increase in cyanobacterial activity. A drastic decline in level-bottom benthic faunas during the FSST of sequence No. 2 is, however, interpreted as a local response to the progradation of a delta complex (the Burgsvik Sandstone). Biological carbonate production replenishes rapidly within the TST of sequence No. 3, succeeding a thin LST dominated by reworked siliciclastics and chemically precipitated carbonates (ooids). The detailed relationship between the CIE and sea-level change presented herein is not fully consistent with previous reports on the CIEs associated with the lower Silurian Ireviken and Mulde events, respectively. Based on our facies analysis and sequence stratigraphical interpretation, two main mechanisms are suggested as responsible for the Late Ludfordian CIE: (1) a change in the riverine C-weathering flux towards the ¹³C end member following glacio-eustatically induced subaerial exposure of carbonate platforms throughout the tropics, and, (2) increased photosynthetic activity by benthic cyanobacteria exaggerating the δ¹³C values of precipitated carbonates.