Primnoa Resedaeformis Research Articles

Geological features supporting deep-sea coral habitat in Atlantic Canada

Geological features supporting cold-water coral habitat in Atlantic Canada are reviewed and exemplified using qualitative field observations from the Scotian margin and Southwest Grand Banks, Newfoundland, in the context of regional geology of the Atlantic Canadian continental margin. Coral habitats are concentrated in areas of shelf-crossing troughs and trough-mouth fans associated with glacial ice streams. Most habitat types are supported by glacial or glaciomarine depositional features, although some are supported by erosional features, probably related to subglacial meltwater erosion. Shelf-break and upper continental slope moraine deposits, subject to strong currents, form current-swept cobble-boulder pavements, forming the principal habitats for the large gorgonian corals Primnoa resedaeformis and Paragorgia arborea. At greater depths, upper slope till tongues are upper continental slope gravelly mud deposits, exposed at the surface as isolated cobbles and boulders in a muddy sand matrix. Common skeletal corals in these environments include the large long-lived gorgonian Keratoisis ornata, the smaller gorgonian Acanthogorgia armata, and, in muddy sand between rocks, the small gorgonian Acanella arbuscula. Erosional environments including friable Tertiary mudstones and semi-consolidated Quaternary sediments, colonized by large gorgonians and other corals, were observed in The Gully, the Stone Fence, and the Southwest Grand Banks. Weak bedrock strength may limit the size of large gorgonian coral colonies. Authigenic carbonate crusts, possibly related to cold seeps, may be locally important in supporting coral growth. Predictive models of coral distribution should consider Quaternary and surficial geology.

Patterns in abundance and size of two deep-water gorgonian octocorals, in relation to depth and substrate features off Nova Scotia

Deep-water corals form unique ecosystems, yet very little is known about factors that regulate their distribution and growth. The abundance and size of two deep-water gorgonian coral species, Paragorgia arborea and Primnoa resedaeformis, and their relationship with depth and substratum cover, were investigated at Northeast Channel, off Nova Scotia, in July 2006. This is the first study to measure abundance and size of these two coral species at depths >500 m in the Canadian Atlantic region. A total of 5 transects between 500 and 1000 m depth were examined using video collected by the remotely operated vehicle ROPOS. Abundance of both species was patchy, but higher at these deeper depths than at <500 m. Abundance generally declined with depth, and was moderately correlated with cover of hard substratum (cobble, boulder, bedrock). These relationships were stronger and less variable for P. resedaeformis than for P. arborea, suggesting that factors such as topographic relief may play an additional role in regulating distributions of P. arborea. Maximum colony height was 125 and 240 cm for P. resedaeformis and P. arborea, respectively, and much greater than recorded for depths <500 m. Overall, colony height and depth relationships were strong for both species, but variable among transects. P. resedaeformis showed a negative relationship with depth, while the opposite was observed for P. arborea, suggesting that the two species are affected differently by factors that vary with depth (e.g. temperature, fishing disturbance). Relationships between colony size and size of attachment stone were stronger for P. arborea, especially for overturned colonies, than for P. resedaeformis, suggesting that availability of suitably coarse substrate may be more important for the long-term persistence of P. arborea colonies.

Ages and growth rates of some deep-sea gorgonian and antipatharian corals of Newfoundland and Labrador

Using bomb-14C dating and growth ring counting methods, we calculate life spans and growth rates of six species of deep-sea corals collected at depths of between 400 and 900 m from the continental slope of Newfoundland and Labrador. Bamboo corals ( Acanella arbuscula , Keratoisis ornata ) and antipatharians ( Stauropathes arctica ) secrete concentric growth rings in their axial skeletons, which were proven to form annually for K. ornata and S. arctica. These species had the lowest radial growth rates of 33 ± 11 to 75 ± 11 μm·year–1. Primnoa resedaeformis and Paramuricea spp. had higher radial growth rates of 83 ± 6 to 215 ± 37 μm·year–1. Paragorgia arborea had the highest radial growth rate of 830 ± 120 μm·year–1. Axial growth rates ranged from 0.56 ± 0.05 cm·year–1 for a specimen of Paramuricea spp. to 2.61 ± 0.45 cm·year–1 for a specimen of Primnoa resedaeformis . Life spans ranged from 18 ± 4 years for a live-collected P. resedaeformis to 200 ± 30 years for a subfossil specimen of K. ornata. Because all of the corals were from heavily fished areas, it is likely that age distributions are biased towards smaller and younger colonies. Recovery of deep-sea corals from fishing-induced damage will take decades to centuries.

Stable C and N isotopic composition of cold-water corals from the Newfoundland and Labrador continental slope: Examination of trophic, depth and spatial effects

With the aim of understanding of the trophic ecology of cold-water corals, this paper explores the tissue δ 13C and δ 15N values of 11 ‘coral’ species (8 alcyonacean, 1 antipatharian, 1 pennatulacean, 1 scleractinian) collected along the Newfoundland and Labrador continental slope. Isotopic results delimit species along continua of trophic level and food lability. With an isotopic signature similar to macrozooplankton, Paragorgia arborea occupies the lowest trophic level and most likely feeds on fresh phytodetritus. Primnoa resedaeformis occupies a slightly higher trophic level, likely supplementing its diet with microzooplankton. Bathypathes arctica, Pennatulacea and other alcyonaceans ( Acanella arbuscula, Acanthogorgia armata, Anthomastus grandiflorus, Duva florida, Keratoisis ornata, Paramuricea sp.) had higher δ 13C and δ 15N values, suggesting these species feed at higher trophic levels and on a greater proportion of more degraded POM. Flabellum alabastrum had an isotopic signature similar to that of snow crab, indicating a primarily carnivorous diet. Isotopic composition did not vary significantly over a depth gradient of 50–1400 m. Coral δ 13C increased slightly (<1‰) from the Hudson Strait to the southern Grand Banks, but δ 15N did not. By modulating the availability and quality of suspended foods, substrate likely exerts a primary influence on the feeding habits of cold-water corals.

Late Holocene radiocarbon and aspartic acid racemization dating of deep-sea octocorals

Primnoa resedaeformis is a deep-sea gorgonian coral with a two-part skeleton of calcite and gorgonin (a fibrillar protein), potentially containing long-term records of valuable paleo-environmental information. For various reasons, both radiocarbon and U/Th dating of these corals is problematic over the last few centuries. This paper explores aspartic acid racemization dating of the gorgonin fraction in modern and fossil specimens collected from the NW Atlantic Ocean. Radiocarbon dating of the fossil specimen indicates a lifespan of 700 ± 100 years, the longest yet documented for any octocoral. Gorgonin amino acid compositions were identical in the fossil and modern specimens, indicating resistance to organic diagenesis. Similar to bone collagen, the fibrillar protein of gorgonin may impose conformational constraints on the racemization of Asp at low temperatures. The rate of racemization of aspartic acid ( d/ l-Asp) was similar to previously published results from an 1800 year old anemone ( Gerardia). The age equation was: age (years BP 2000 AD) = [( d/ l − 0.020 (±.002))/.0011 (±.0001)] 2 ( r 2 = 0.97, p < .001). The error in an age estimate calculated by d/ l-Asp was marginally better than that for 14C dating over the most recent 50–200 years, although the dating error may be improved by inclusion of more samples over a broader time range. These results suggest that d/ l-Asp dating may be useful in augmenting 14C dating in cases where 14C calibrations yield two or more intercept ages, or in screening samples for further 14C or U/Th dating.

Megafauna associated with assemblages of deep-water gorgonian corals in Northeast Channel, off Nova Scotia, Canada

The distribution and abundance of benthic megafauna in areas known to be inhabited by dense gorgonian coral assemblages were examined at Northeast Channel, off Nova Scotia, Canada, in August 2001. Using a remotely operated vehicle, 1–5 video transects during each of 1–2 dives at each of four sites (Rips, Middle Canyon, Hell Hole West and Hell Hole East) were conducted. The relationships in the structure of biological assemblages at three spatial scales: within transects (10s of metres); between dive locations (100s of metres); and among sites (10s to 100s of kilometres) were explored. The most abundant epibenthic taxa included the gorgonians Primnoa resedaeformis and Paragorgia arborea, several suspension feeders (Actinauge verrilli, Bolocera tudiae, an unidentified anemone and encrusting sponge, Ophiacantha abyssicola), the deposit feeder Porania pulvillus insignus and the predatory Solaster endeca. The basket star Gorgonocephalus arcticus was present only on colonies of Paragorgia arborea. Despite large variability in abundance and assemblage composition among transects and dive locations, clear patterns were observed among sites. Mean abundance of most cnidarians and echinoderms was greatest at Hell Hole West. No gorgonians were found at Hell Hole East. The encrusting sponge was most abundant at the Rips and least abundant at Hell Hole East. Cluster analysis and multidimensional scaling (MDS) indicated that, when abundance was averaged across transects for each dive, the megafaunal assemblages fall into groups of dives that separated by site. These differences among sites are most likely related to variability in the physical environment. The epibenthic megafaunal assemblages were as diverse in the presence as in the absence of gorgonian corals, at least at the abundances that we observed. However, the apparent low recruitment and abundance, combined with small population size make these assemblages particularly vulnerable to perturbations.

Morphology and growth of the deep-water gorgonians Primnoa resedaeformis and Paragorgia arborea

The morphology of the gorgonian corals Paragorgia arborea and Primnoa resedaeformis was studied from video records and colonies collected from different locations in Atlantic Canada, at depths between 200 and 600 m. Growth was studied by relating colony height to age (number of growth rings) in P. resedaeformis, and from a photographic time-series of a P. arborea colony in a Norwegian fjord. The highest P. resedaeformis and P. arborea colonies were 86 and 180 cm, respectively. The height of P. arborea seemed to be restricted by the size of the boulder it was attached to. When the coral exceeds a critical height (approximately twice the stone size), the drag of strong currents can turn the coral and its substrate over. No limiting factors for the height of P. resedaeformis colonies were identified. P. arborea occurred in three colour varieties: red, salmon red, and white. The red and white contributed 41% to the population each, while 18% of the colonies were salmon red. On average the salmon red P. arborea were taller than the red and white. P. arborea colonies >50 cm were mainly concave fan shaped. The orientation of these indicated a near-bottom current pattern similar to what is known from previous current measurements in the area. P. resedaeformis occurred mainly on the up-current side of boulders, but its bushy morphology does not indicate influence by unidirectional current to the same degree as P. arborea. The different height, morphology, and position on boulders of the two species indicate that they utilize different food sources. P. resedaeformis seems to be adapted to a near-bottom environment with turbulent currents, whereas P. arborea utilize uni- or bidirectional currents higher above bottom by developing planar colonies perpendicular to the current. The oldest P. resedaeformis colony was 61 years. The relationship between height and age indicated an average growth of 1.7 cm year−1 for P. resedaeformis. X-ray images of skeletal sections of P. arborea showed clear growth bands with a maximum band width of 1.3 cm. It is not clear what time scales these bands represent, and they could therefore not be used for indicating age. The limited previously reported data on age and growth of P. arborea indicate an average growth rate of 1 cm year−1. This gives an age of about 180 years for the largest colony in this study. The time-series photographs, however, indicated a much higher growth rate (varying between 2 and 6 cm year−1 within the colony), which may be more representative for colonies of an intermediate size.

Radiocarbon evidence for annual growth rings in the deep-sea octocoral Primnoa resedaeformis

The deep-sea gorgonian octocoral Primnoa resedaeformis is distributed throughout the Atlantic and Pacific Oceans at depths of 65-3200 m. It has a two-part skeleton of calcite and gorgonin. Towards the inside of the axial skeleton gorgonin and calcite are deposited in concentric growth rings, similar to tree rings. Colonies were collected from the Northeast Channel (northwest Atlantic Ocean, southwest of Nova Scotia, Canada) from depths of 250-475 m. Radiocarbon was measured in individual rings isolated from sections of each colony, after dissolution of calcite. Each {Delta}{sup 14}C measurement was paired with a ring age determined by three amateur ring counters. The precision of ring counts averaged better than {+-} 2 years. Accurate reconstruction of 20th century bomb-radiocarbon shows that (1) the growth rings are formed annually, (2) the gorgonin is derived from surface particulate organic matter (POM) and (3) useful environmental data are recorded in the organic endoskeletons of deep-sea octocorals. These results support the use of Primnoa resedaeformis as a long-term, high resolution monitor of surface ocean conditions, particularly in temperate and boreal environments where proxy data are lacking.

Crustaceans associated with the deep-water gorgonian corals Paragorgia arborea (L., 1758) and Primnoa resedaeformis (Gunn., 1763)

To explore the crustacean fauna associated with deep-water gorgonian corals, suction samples were taken from colonies of Paragorgia arborea and Primnoa resedaeformis using a Remotely Operated Vehicle. Seven colonies of P. arborea and eight of P. resedaeformis were sampled from 330–500 m depth in the Northeast Channel off Nova Scotia. A total of 17 species were identified as being associated with the corals. The P. arborea fauna was richer than the P. resedaeformis fauna in both abundance and number of species, with 1303 versus 102 individuals and 16 versus seven species, respectively. However, 13 of the species associated with P. arborea were from hydroids attached to the coral. Amphipods dominated the fauna both in abundance and numbers of species and the most common species were Metopa bruzelii, Stenopleustes malmgreni, Proboloides calcarata and Aeginella spinosa. The isopod Munna boecki and the cirripede Ornatoscalpellum stroemii were also quite common. The most strongly associated crustaceans were two parasitic poecilostomatid copepods; these are common also on tropical gorgonians and are most likely obligate associates. The frequently occurring shrimp Pandalus propinquus probably avoids predation by seeking protection among the coral branches. Shrimp counts from video records showed that visual inspection without physically disturbing colonies will generally not reveal the crustaceans hidden in coral colonies. The galatheid Eumunida picta was observed on P. resedaeformis colonies. The fauna of the deep-water gorgonians corresponded to the fauna of tropical shallow-water gorgonians in the numerical dominance of amphipods and parasitic copepods; however, species richness is higher and decapods, which constitute a rich fauna on tropical gorgonians, were only represented by two species.

Distribution of deep-water gorgonian corals in relation to benthic habitat features in the Northeast Channel (Atlantic Canada)

The distribution and abundance of deep-water gorgonian corals were investigated along 52 transects at 183–498 m depth in the Northeast Channel, between Georges Bank and Browns Bank in the northwest Atlantic, using a remotely operated vehicle and a towed video-camera system. Three species (Paragorgia arborea, Primnoa resedaeformis, and Acanthogorgia armata) were observed. Primnoa occurred on 35 transects below 196 m depth, with highest local abundance in stands of 104 colonies per 100 m2. Paragorgia was present on 21 transects deeper than 235 m, with highest local abundance of 49 colonies per 100 m2. Acanthogorgia was observed at only four transects between 231 m and 364 m, with a local maximum abundance of 199 colonies per 100 m2. The maximum abundance averaged for whole transects was 19.2 and 6.2 colonies per 100 m2 for Primnoa and Paragorgia, respectively. The corals were more common in the outer part of the channel along the shelf break and slope than on the shelf in the inner part. All three species showed a patchy distribution with no signs of competitive exclusion at any spatial scale. Transects with high abundance of corals were characterised by depths greater than 400 m, maximum temperatures less than 9.2°C, and a relatively high percentage coverage of cobble and boulder (more than 19% and 6%, respectively). High temperatures probably control the upper depth limit of the corals, and Primnoa seems to tolerate slightly higher temperatures than Paragorgia. Abundance of both species was negatively correlated with average temperature and positively with cobbles. Together, temperature, percentage cobble and salinity accounted for 38% of the variance of Primnoa. The comparable figure for Paragorgia was 15%. The observed distribution indicated that the abundance of coral is controlled by additional factors such as larger-scaled topographic features governing the current regimes and thus also the supply of food and larvae.

Potential climate signals from the deep-sea gorgonian coral Primnoa resedaeformis

The deep-sea gorgonian coral Primnoa resedaeformis has an arborescent skeleton composed of both calcite and a horn-like structural protein called gorgonin. We have investigated potential climate records in corals from Alaska, the eastern seaboard of Canada and the United States, and a Southern Ocean (Pacific sector) seamount. Temperatures at these sites range from 4 to ∼10 °C. δ18O values of the calcite show strong evidence for isotopic disequilibrium. Extraction of δ18O paleotemperatures is therefore not straightforward. Sr/Ca data, analyzed by Secondary Ion Mass Spectrometry (SIMS), suggest that temperature might be a control on calcite Sr/Ca in Primnoa resedaeformis, but that growth-related kinetic effects could also be important. Based on previous 14C, δ13C and δ15N measurements, it has been suggested that particulate organic matter (POM) from the surface is an important carbon source to the polyps and the gorgonin fraction of the Primnoa skeleton. δ15N and δ13C of polyps and gorgonin show similar regional differences to δ15N and δ13C of surface POM. Polyps and contemporaneous gorgonin correlate strongly for both δ13C and δ15N. The influence of nutrient isotopic composition and climate and productivity variations on the isotopic composition of surface POM may therefore be recorded in gorgonin layers. These corals have very long lifespans (several centuries). The potential exists, therefore, to obtain extended records of surface productivity, deep ocean temperature and chemistry of value to climatologists and fisheries managers.

Age, growth and radiometric age validation of a deep-sea, habitat-forming gorgonian (Primnoa resedaeformis) from the Gulf of Alaska

Sustainable fisheries require (1) viable stock populations with appropriate harvest limits and (2) appropriate habitat for fish to survive, forage, seek refuge, grow and reproduce. Some deep-water habitats, such as those formed by deep-water stands of coral, may be vulnerable to fishing disturbance. The rate at which habitat can be restored is a critical aspect of fishery management. The purpose of this study was to characterize growth rates for a habitatforming deep-sea coral. Two nearly complete colonies of red tree coral (Primnoa resedaeformis) collected from waters off southeast Alaska were used for an analysis of age and growth characteristics. CAT scans revealed that colonies consisted of multiple settlement events, where older basal structures provided for settlement of new colonies. The decay of 210 Pb over the length of the colony was used to validate age estimates from growth ring counts. Age estimates were over 100 yr for sections near the heavily calcified base. Based on validated growth ring counts, growth of red tree coral ranged from 1.60 to 2.32 cm per year in height and was approximately 0.36 mm per year in diameter. These growth rates suggest that the fishery habitat created by red tree coral is extremely vulnerable to bottom fishing activities and may take over 100 years to recover.

Lifespans and growth patterns of two deep-sea corals: Primnoa resedaeformis and Desmophyllum cristagalli

A subfossil fragment of the deep-sea gorgonian coral Primnoa resedaeformis was 14C AMS dated along a radial growth transect. Dates ranged from 2600±50 at the outside, to 2920±60 14C years BP near the interior, suggesting an age of <300 years. The avergae radial growth was approximately 0.044 mm.yr−1. Based on comparisons with live-collected specimens, we estimate the entire colony may have been about 0.5–0.75 m tall, with a linear tip extension rate of 1.5–2.5 mm.yr−1. Towards the centre of the main stem, the coral skeleton is composed of alternating couplets, 200 μm in width, of gorgonin (a horn-like organic skeletal protein) and calcite. We believe these couplets are annual. Within this larger scale of banding are finer couplets of gorgonin and calcite, with frequencies suggesting lunar monthly periodicity. Both scales of banding may reflect fluctuations in food supply from sinking POM, or from tidally-resuspended bottom POM, along with benthic consumers. Outer skeletal growth is predominantly massive calcite with intermittent gorgonin layers. If carbonate precipitation in this zone were continuous, approximately 25 μm radial growth would be deposited every year. The Scleractinian Desmophyllum cristagalli lives to <200 years, and has rates of linear extension of 0.5–1.0 mm.yr−1. The skeletons show growth bands approximately 10 μm wide, which may be annual. Due to tissue extension and retraction in life, parts of the skeleton may be overgrown, or suffer dissolution. Although we have shown in previous publications that sea water temperatures may be obtained from analysis of this coral, periods of skeletal dissolution, coupled with isotopic disequilibrium, will make obtaining long climatic records extremely difficult.

Post-Glacial Interchange between Pacific and Atlantic Deeper Living Boreal Faunas?

Investigators of the animal world of northern waters cannot without interest pass by a small group of species, which, geographically, provoke our curiosity by reason of their discontinuously circumterrester occurrence in the boreal region, living both in Atlantic and Pacific waters. We may here e.g. name the hydroid, Bonneviella grandis (Allman), the hydrocorals, Stylaster (Eu-Stylaster) gemmascens (Esper) and Stylaster (Allopora) norvegicus (Gunnerus), and the octocorals, Paragorgia arborea (Linne) and Primnoa resedaeformis (Gunnerus). None of them is able to live in the icy cold layers of the Arctic seas, nor can they, on the other hand, endure the higher temperatures of tropical and sub-tropical waters. The species do not belong to shallow water; nor can they penetrate into deep-sea proper. According to our present knowledge their bathymetrical ranges are: