Commensal symbiosis between agglutinated polychaetes and sulfate‐reducing bacteria

Pendant bioconstructions occur within submerged caves in the Plemmirio Marine Protected Area in SE Sicily, Italy. These rigid structures, here termed biostalactites, were synsedimentarily lithified by clotted‐peloidal microbial carbonate that has a high bacterial lipid biomarker content with abundant compounds derived from sulfate‐reducing bacteria. The main framework builders are polychaete serpulid worms, mainly Protula with subordinate Semivermilia and Josephella. These polychaetes have lamellar and/or fibrillar wall structure. In contrast, small agglutinated terebellid tubes, which are a minor component of the biostalactites, are discontinuous and irregular with a peloidal micritic microfabric. The peloids, formed by bacterial sulfate reduction, appear to have been utilized by terebellids to construct tubes in an environment where other particulate sediment is scarce. We suggest that the bacteria obtained food from the worms in the form of fecal material and/or from the decaying tissue of surrounding organisms and that the worms obtained peloidal micrite with which to construct their tubes, either as grains and/or as tube encompassing biofilm. Peloidal worm tubes have rarely been reported in the recent but closely resemble examples in the geological record that extend back at least to the early Carboniferous. This suggests a long‐lived commensal relationship between some polychaete worms and heterotrophic, especially sulfate‐reducing, bacteria.     

A review of the status and decline in abundance of the Irish Hare (Lepus timidus hibernicus) in Northern Ireland

The Northern Ireland Hare Survey documented the distribution of the Irish Hare (Lepus timidus hibernicus). Historical game bag records and other, more contemporary, records of hare distribution were examined. These data indicate how numbers of L. t. hibernicus may have changed over the last 140 years. The results of the Northern Ireland Hare Survey suggested that L. t. hibernicus was widespread throughout Northern Ireland. Current average densities are no more than 0.65 hares/km². Game bag records indicate that hare densities may have been much higher in the past, with a maximum of 138 hares/km² recorded on Crom Estate, Co. Fermanagh, in 1864. Evidence from hare distribution recorded during the Northern Ireland Rabbit Survey indicates that hare numbers declined between 1984 and 1994. Evidence from all sources suggests that L. t. hibernicus has declined in abundance substantially, with present total population estimates for Northern Ireland ranging from 8250 to 21 000 individuals. Flushing data indicate that rushes and hedgerows are important diurnal resting areas for hares. While the principal reason for the decline in numbers of L. t. hibernicus in Northern Ireland is not clear, more species‐rich pasture and provision of areas of cover, such as rushes, may arrest further declines, or indeed promote numbers of hares, particularly in lowland areas.     

The effects of symbiotic crabs on the pumping activity and growth rates of Chaetopterus variopedatus

This study investigates how the presence of symbiotic crabs (Pinnixa chaetopterana or Polyonyx gibbesi) in the tubes of the polychaete Chaetopterus variopedatus affects the worms' pumping activity and growth rates under laboratory and field conditions. In the field, worms whose tubes are inhabited by Pinnixa beat their fan segments significantly more frequently than do worms hosting Polyonyx, but other measures of pumping activity do not differ according to symbiont species. In the lab, worms tend to move water through their tubes at higher rates when crabs are present. In 7-month laboratory experiments, growth rates of worms hosting either species of crab did not differ from growth rates of worms without crab symbionts. Although worms hosting Polyonyx are, on average, significantly larger than worms hosting Pinnixa, this appears to be due to competition between the crab species for hosts and not due to differential effects on host growth. Unlike the crabs in this study, pea crab species inhabiting bivalves are known to have strong deleterious effects on host growth and reproduction, suggesting that the evolution of virulence in symbiotic interactions is dependent upon specific ecological context.     

Mineralization of Alvinella polychaete tubes at hydrothermal vents

Alvinellid polychaete worms form multilayered organic tubes in the hottest and most rapidly growing areas of deep‐sea hydrothermal vent chimneys. Over short periods of time, these tubes can become entirely mineralized within this environment. Documenting the nature of this process in terms of the stages of mineralization, as well as the mineral textures and end products that result, is essential for our understanding of the fossilization of polychaetes at hydrothermal vents. Here, we report in detail the full mineralization of Alvinella spp. tubes collected from the East Pacific Rise, determined through the use of a wide range of imaging and analytical techniques. We propose a new model for tube mineralization, whereby mineralization begins as templating of tube layer and sublayer surfaces and results in fully mineralized tubes comprised of multiple concentric, colloform, pyrite bands. Silica appeared to preserve organic tube layers in some samples. Fine‐scale features such as protein fibres, extracellular polymeric substances and two types of filamentous microbial colonies were also found to be well preserved within a subset of the tubes. The fully mineralized Alvinella spp. tubes do not closely resemble known ancient hydrothermal vent tube fossils, corroborating molecular evidence suggesting that the alvinellids are a relatively recent polychaete lineage. We also compare pyrite and silica preservation of organic tissues within hydrothermal vents to soft tissue preservation in sediments and hot springs.     

Ecology and status of Erica vagans in County Fermanagh, Ireland

The colony of Erica vagans in County Fermanagh, Northern Ireland was surveyed and soil and species-association analyses were carried out. The species is found in a base-rich flush, were it survives due to reduced competition from calcifuge species and is associated with other species that tolerate the relatively calcareous environment. The colony is uniquely uniform in flower colour and does not appear to be able to reproduce sexually at this time. The species probably arrived in Ireland in postglacial times and has since becomegeneticallydistinct.fi. vagans does not appear to be a recent introduction, though man may have been involved in determining its present distribution and that of the other so-called "Hiberno-Lusitanian"Ericaceae. While considerable doubt remains concerning the origin and age of the colony, the species cannot be regarded as native.     

Chaetae and mechanical function: tools no Metazoan class should be without

To address the functional contributions of capillary chaetae in the maldanid polychaete Clymenella torquata, we compared irrigation efficiency and tube structure for animals with intact and trimmed capillary chaetae. We measured pumping rates for worms before and after they were anaesthetized and subjected either to capillary trimming or mock trimming, i.e. handling without trimming. Worms with trimmed chaetae were significantly less effective at moving water through their tubes than those with intact chaetae. There were no significant differences in the ability of control worms to move water within their tubes. No significant changes in rates of peristalsis were observed among experimental or control groups. These data strongly suggest that body musculature and capillary chaetae work in concert to hold worms in position within tubes during peristaltic pumping. When chaetae are shortened, the body musculature must contract to a greater degree, increasing the functional diameter of the worm to achieve the necessary traction with the tube wall, resulting in less efficient irrigation. We also compared the inner diameters of original field tubes to tubes built by control worms or worms after capillary trimming. The inner diameters of new tubes built by worms with shortened chaetae were larger than their original tubes, while those of both control groups were not. One possible explanation is that the chaetae have a sensory role and shortened chaetae send the false message that the nascent tube walls are farther away than they are, the body contracts in compensation and the tube is widened, however this idea has not been tested.     

HOLDING ON BY THEIR HOOKS: ANCHORS FOR WORMS

We examined the hooked setae of a dominant group of tube-dwellers, the polychaete annelids, and found a pattern of setation that is predictable by tube type, exclusive of worm taxon or orientation; we also demonstrated the mechanical significance of these hooked setae. When tube-dwelling worms belonging to different lineages are pressurized, they resist differentially as a function of the direction in which hooks face. The results are consistent with the hypothesis that hooks are used primarily to resist removal of worms from their tubes, are polyphyletic in origin, and are active agents of resistance.     

Polychaete worms--a vector for white spot syndrome virus (WSSV)

The present work provides the first evidence of polychaete worms as passive vectors of white spot syndrome virus (WSSV) in the transmission of white spot disease to Penaeus monodon broodstocks. The study was based on live polychaete worms, Marphysa spp., obtained from worm suppliers/worm fishers as well as samples collected from 8 stations on the northern coast of Tamilnadu (India). Tiger shrimp Penaeus monodon broodstock with undeveloped ovaries were experimentally infected with WSSV by feeding with polychaete worms exposed to WSSV. Fifty percent of polychaete worms obtained from worm suppliers were found to be WSSV positive by 2-step PCR, indicating high prevalence of WSSV in the live polychaetes used as broodstock feed by hatcheries in this area. Of 8 stations surveyed, 5 had WSSV positive worms with prevalence ranging from 16.7 to 75%. Polychaetes collected from areas near shrimp farms showed a higher level of contamination. Laboratory challenge experiments confirmed the field observations, and > 60% of worms exposed to WSSV inoculum were proved to be WSSV positive after a 7 d exposure. It was also confirmed that P. monodon broodstock could be infected with WSSV by feeding on WSSV contaminated polychaete worms. Though the present study indicates only a low level infectivity in wild polychaetes, laboratory experiments clearly indicated the possibility of WSSV transfer from the live feed to shrimp broodstock, suggesting that polychaete worms could play a role in the epizootiology of WSSV.     

Paleozoic shell-Boring Annelids and their Trace Fossils

Polychaete shell-borings are widespread but not often commonin Paleozoic strata of most continents, occurring in pelecypod,gastropod, cephalopod, coral, stromatoporoid, crinoid, brachiopod,ectoproct, and calcareous algal skeletons of Ordovician to Permianage. Four genera are known: Vermiforichnus (Spionidae), Ordovicianto Permian; Caulostrepsis (Spionidae) , Devonian to Oligocene;Myzoslomites (Myzostomidae), Ordovician to Jurassic: and possiblyConchotrema, Devonian to Permian. The Silurian Citonia siphois a worm-boring, not a calcareous worm tube, referable to Vermiforichnus. An exceedingly well preserved Devonian fossilized polychaete,Vermiforafacta rollhisi, with complex setigerous parapodia,dorsal cirri, peristomial cirri, prostomium, and tentacle-likepalps, occurs entombed in an agglutinated tube-lined worm-boring(Vermiforichnus) . It may have been a shell-borer consideringits many similarities to the spionid, Polydora, which apparentlyecologically replaced it. Like Polydora, Vermiforafacta probably filtered food and sedimentfrom sea water and constructed agglutinated tubes which linedits borings. In bivalves, borings parallel valves and are usuallyperpendicular to commissures; in massive skeletons, they radiateoutward and laterally. Distorted later growth and "blisters"indicate that the host was often alive. Commensal to pathogenicparasitism existed and worms benefited from their hosts's feedingcurrents and protective shells. One polychaete nestler is reported.Life positions of hosts are sometimes indicated.     

Feeding, growth, and fecundity of Abarenicola pacifica in relation to sediment organic concentration

For marine deposit-feeding invertebrates, the distribution of species with different life history strategies has long been known to be correlated with sediment organic concentration. Large populations of opportunistic species are found in sediments with enriched organic concentration, while equilibrium species populate low organic concentration sediments. Differences in both behavioral (e.g. feeding rate) and physiological (e.g. growth rate, reproductive output) adaptations determine the ability of species to establish populations in different environments. By systematically documenting differences in the way these factors vary as sediment organic concentration varies for both opportunistic and equilibrium species, we can better understand the mechanisms underlying this correlation between sediment organic concentration and species distributions. Here, we present the results of experiments examining the interactions among food concentration, feeding rate, growth rate, and reproductive output (measured as egg number and size) for the equilibrium species Abarenicola pacifica. A. pacifica is a large, long-lived, iteroparous, sub-surface deposit-feeding polychaete. Individual worms were reared throughout most of one generation in sediments differing only in the concentration of organic matter. Juveniles (<20 mg AFDW) had higher feeding rates and growth rates in sediments of higher organic concentration throughout the range tested. These results are consistent with the predictions from optimal foraging theory. As worms grew, however, these patterns changed. Once worms reached a mean body size of approximately 50 mg AFDW, feeding rate was greater on sediments of lower organic concentration (although it took worms in the sediments with lower organic concentration longer to reach this size). Differences in growth rates among treatments decreased as worms grew. For worms >100 mg AFDW, growth rates were uniformly low ( approximately 1%/day) on all sediments, but the early advantage obtained by worms in the high organic treatments resulted in much greater body sizes after 200 days. Worms had higher tissue triacylglyceride concentrations and produced more eggs (independent of worm size) as sediment organic concentration increased. We conclude that A. pacifica alters its feeding rate in response to variations in food resources in such a way as to maximize its energy intake and thereby maximize fitness. Future studies should investigate whether opportunistic species (as well as other equilibrium species) also have this ability.     

Oxygen consumption and feeding rates of the sabellid polychaete, Myxicola infundibulum (Renier)

1. Rates of respiration and feeding were measured for the sabellid polychaete, Myxicola infundibulum. 2. Basal rates of respiration were 1.5 times lower than routine rates. 3. Clearance and irrigation rates were independent of cell type when the worms were fed on monocultures of algae; however, preferential selection was seen when the algal species were offered simultaneously. 4. Calculations indicate that this species can meet its energetic needs at rather low cell densities. 5. It is suggested that these low energy requirements, coupled with the capacity for high pumping/clearing rates may be of adaptive significance in reducing competition for food resources.     

Attached vermiform gastropods in Carboniferous marginal marine stromatolites and biostromes

Tubiform fossils conventionally referred to Serpula cf. advena Salter and species of Spirorbis Lamarck from the British Lower Limestone Shales and Border Group (Lower Carboniferous) are re-examined. They occur in peritidal carbonate environments of schizohaline aspect. These fossils superficially resemble calcareous polychaete tubes but have skeletal characters, including molluscan wall structure, numerous internal septa, and protoconch, which indicate that they represent a new group of substrate-attached, disjunctly coiled gastropods. They resemble archaeogastropods in internal morphology of the skeleton but show parallels in external form and occurrence with the extant Vermetidae. There are two principal modes of occurrence: (1) erect tubes forming intertidal biostromes associated with non-skeletal algal laminites, and (2) prostrate discoidal tubes encrusting subtidal skeletal stromatolites or occasionally forming larger irregular bioherms. These biostromes and bioherms are comparable in structure to Recent vermetid reef developments.     

First evidence of maternal transmission of algal endosymbionts at an oocyte stage in a triploblastic host, with observations on reproduction in Waminoa brickneri (Acoelomorpha)

Abstract. Examination of sexual reproduction in a symbiotic acoelomorph worm, Waminoa brickneri from Eilat (Red Sea), presents the first definitive evidence for maternal transmission of dinoflagellate algal symbionts in a triploblastic organism. Sexually mature worms were removed from the stony coral Plesiastrea laxa and raised in the laboratory. Eggs were detected 18 d after the collection of the worms and hatched 4 d later. Histological sections performed on sexually mature worms showed an ovary with oocytes containing two distinct types of algal endosymbionts within their ooplasm. Transmission electron microscopy corroborated the presence of algal symbionts within the developing embryos. Our findings regarding maternal transmission of symbionts shed new light on the diversity of modes of algal symbiont acquisition known in triploblastic organisms.     

Textures and traction: how tube‐dwelling polychaetes get a leg up

By controlling the traction between its body and the tube wall, a tube‐dwelling polychaete can move efficiently from one end of its tube to the other, brace its body during normal functions (e.g., ventilation and feeding), and anchor within its tube avoiding removal by predators. To examine the potential physical interaction between worms and the tubes they live in, scanning electron microscopy was used to reveal and quantify the morphology of worm bodies and the tubes they produce for species representing 13 families of tube‐dwelling polychaetes. In the tubes of most species there were macroscopic or nearly macroscopic (~10 μm–1 mm) bumps or ridges that protruded slightly into the lumen of the tube; these could provide purchase as a worm moves or anchors. At this scale (~10 μm‐1 mm), the surfaces of the chaetal heads that interact with the tube wall were typically small enough to fit within spaces between these bumps (created by the inward projection of exogenous materials incorporated into the tube wall) or ridges (made by secretions on the interior surface of the tube). At a finer scale (0.01–10 μm), there was a second overlap in size, usually between the dentition on the surfaces of chaetae that interact with the tube walls and the texture provided by the secreted strands or microscopic inclusions of the inner linings. These linings had a surprising diversity of micro‐textures. The most common micro‐texture was a “fabric” of secreted threads, but there were also orderly micro‐ridges, wrinkles, and rugose surfaces provided by microorganisms incorporated into the inner tube lining. Understanding the fine structures of tubes in conjunction with the morphologies of the worms that build them gives insight into how tubes are constructed and how worms live within them.     

Palp growth,regeneration, and longevity of the obligate hermit crab symbiont Dipolydora commensalis (Annelida: Spionidae)

Abstract. The polychaete Dipolydora commensalis is an obligate symbiont of hermit crabs and produces a burrow along the columella of the gastropod shells they inhabit. Adults of D. commensalis have short palps that they use to feed on particles dropped or brought in by the respiratory currents of hosts. To determine whether hermit crabs influence palp length, specimens of D. commensalis were isolated in glass capillary tubes and the growth of palps was measured over a 3‐week period. Palp length was also measured in worms isolated in gastropod shells with or without hermit crabs for 2 weeks. In addition, to determine whether adults of D. commensalis have regeneration capabilities like those of free‐living relatives, worms were cut at the fifth or 15th setiger and then monitored for 35 d. Worms extracted from shells and placed into capillary tubes had initial palp lengths of 1.0±0.4 mm (n=17); after isolation, palps were 40% longer (1.4±0.4 mm, n=17). Worms in gastropod shells with hermit crabs had an average palp length of 0.9±0.4 mm (n=31), whereas worms in shells without hermit crabs had palps that were 33% longer (1.2±0.5 mm, n=40). Adults of D. commensalis are capable of regeneration; 35 d after ablation at setigers 5 or 15, the average number of anterior setigers regenerated was 5 (n=15) and 9±1.3 (n=13), respectively. The average number of posterior setigers regenerated from the 15 setiger anterior fragments was 11±6 (n=10). The findings suggest that the palps (and sometimes anterior ends) of the worms are exposed during feeding and are cut during movement of the hermit crab. In the laboratory worms can live for >4 years, considerably longer than the functional life span of most gastropod shells inhabited by hermit crabs.     

Palynological correlation of the upper devonian and lower carboniferous in central Ireland

Miospore assemblages assigned to the PL, NV, CM and Pu Zones have been recorded from a number of boreholes which penetrated Upper Devonian and Lower Carboniferous (Tournaisian and early Visean) rocks in central Ireland. The palynomorphs occur in both marine and non-marine successions, permitting correlation between these two facies. The boundary between the PC and CM Zones has been located within three sections. The palynological data suggest that the base of the marine sequence is of mid-Tournaisian (Tn2) age at Ballyvergin in the south of the area described and of uppermost Tournaisian (Tn3) or earliest Visean age around Clew Bay in the northwest.     

Impacts of forestry planting on primary production in upland lakes from north‐west Ireland

Planted forests are increasing in many upland regions worldwide, but knowledge about their potential effects on algal communities of catchment lakes is relatively unknown. Here, the effects of afforestation were investigated using palaeolimnology at six upland lake sites in the north‐west of Ireland subject to different extents of forest plantation cover (4–64% of catchment area). ²¹⁰Pb‐dated sediment cores were analysed for carotenoid pigments from algae, stable isotopes of bulk carbon (δ¹³C) and nitrogen (δ¹⁵N), and C/N ratios. In lakes with >50% of their catchment area covered by plantations, there were two‐ to sixfold increases in pigments from cryptophytes (alloxanthin) and significant but lower increases (39–116%) in those from colonial cyanobacteria (canthaxanthin), but no response from biomarkers of total algal abundance (β‐carotene). In contrast, lakes in catchments with <20% afforestation exhibited no consistent response to forestry practices, although all lakes exhibited fluctuations in pigments and geochemical variables due to peat cutting and upland grazing prior to forest plantation. Taken together, patterns suggest that increases in cyanobacteria and cryptophyte abundance reflect a combination of mineral and nutrient enrichment associated with forest fertilization and organic matter influx which may have facilitated growth of mixotrophic taxa. This study demonstrates that planted forests can alter the abundance and community structure of algae in upland humic lakes of Ireland and Northern Ireland, despite long histories of prior catchment disturbance.     

Ordovician reefs of the Ural Mountains,Russia: A review

Summary The Upper Ordovician reefs of the Urals were formed at a subsiding shelf-margin during an early Late Ashgillian (Sur’ya time interval) regressive phase. Reefs of this age were studied in detail from the western slope of the Northern, the Subpolar and the Polar Urals with respect to lithofacies, biotic composition and paleogeographical patterns. The thickness of the reefs varies between 100 and 500 m. The backreef areas are characterized by lagoons with increased salinity and sabkha development. Microbial associations and a diverse algal flora (Cyanophyta, green and red algae and alga incertae sedis) are the main constituents of reefal boundstones. Tabulate and rugose corals, heliolitids, calcareous sponge-like fossils, bryozoans and problematic hydroids were also part of the reef communities. Each reef exhibits a characteristic framework-building association. Reef development was terminated by a rapid and abrupt sea-level rise at the end of the middle Upper Ashgillian connected with the global Late Ordovician glaciation.     

A species distribution model for pine marten (<Emphasis Type="Italic">Martes martes</Emphasis>) in the least forested region of Europe

Species distribution modelling is a useful technique that provides data on factors that can influence a species range, identify high suitability areas and model future scenarios. The pine marten (Martes martes) has undergone major historical declines in abundance and distribution in Northern Ireland, similar to that which has occurred throughout its range. Currently, the species is in a phase of range expansion in Northern Ireland, in what is the least forested landscape in Europe. To assess the suitability of this environment for pine marten re-establishment, presence only distribution data combined with landcover data at a 10-km scale were used in a species distribution modelling study using Maxent. The results indicated that approximately 32% (4500 km²) of the land area of Northern Ireland had a high probability of pine marten occurrence. Pine marten distribution was positively associated with the extent of conifer forest landcover types, which also had the highest single attribute contribution to the model. Landcover types that were negatively associated with pine marten distribution included the extent of open, dwarf and urban areas.