Red Tree Coral Research Articles

Shallow-emerged coral may warn of deep-sea coral response to thermal stress

In the Gulf of Alaska, commercially harvested fish species utilize habitats dominated by red tree corals (Primnoa pacifica) for shelter, feeding, and nurseries, but recent studies hint that environmental conditions may be interrupting the reproductive lifecycle of the corals. The North Pacific has experienced persistent and extreme thermal variability in recent years and this pattern is predicted to continue in coming decades. Recent discovery of deep-water emerged coral populations in Southeast Alaska fjords provided opportunity for detailed life-history studies and comparison to corals in managed habitats on the continental shelf. Here we show that sperm from deep colonies develops completely, but in shallow colonies, sperm development is prematurely halted, likely preventing successful production of larvae. We hypothesize that the divergence is due to differing temperature regimes presently experienced by the corals. Compared to deep populations below the thermocline, shallow populations experience much greater seasonal thermal variability and annual pulses of suspected near-lethal temperatures that appear to interrupt the production of viable gametes. The unique opportunity to comprehensively study emerged populations presently affected by thermal stress provides advance warning of the possible fate of deep corals in the Gulf of Alaska that will soon experience similar ocean conditions.

Understanding growth and age of red tree corals (Primnoa pacifica) in the North Pacific Ocean.

Massive, long-lived deep-sea red tree corals (Primnoa pacifica) form a solid, layered axis comprised of calcite and gorgonin skeleton. They are abundant on the outer continental shelf and upper slope of the Northeast Pacific, providing habitat for fish and invertebrates. Yet, their large size and arborescent morphology makes them susceptible to disturbance from fishing activities. A better understanding of their growth patterns will facilitate in-situ estimates of population age structure and biomass. Here, we evaluated relationships between ages, growth rates, gross morphological characteristics, and banding patterns in 11 colonies collected from depths of ~141–335 m off the Alaskan coast. These corals ranged in age from 12 to 80 years old. They grew faster radially (0.33–0.74 mm year-1) and axially (2.41–6.39 cm year-1) than in previously measured older colonies, suggesting that growth in P. pacifica declines slowly with age, and that basal diameter and axial height eventually plateau. However, since coral morphology correlated with age in younger colonies (< century), we developed an in-situ age estimation technique for corals from the Northeast Pacific Ocean providing a non-invasive method for evaluating coral age without removing colonies from the population. Furthermore, we determined that annual bands provided the most accurate means for determining coral age in live-collected corals, relative to radiometric dating. Taken together, this work provides insight into P. pacifica growth patterns to inform coastal managers about the demographics of this ecologically important species. With this new ability to estimate the age of red tree corals in-situ, we can readily determine the age-class structure and consequently, the maturity status of thickets, using non-invasive video survey techniques when coupled with mensuration systems such as lasers or stereo-cameras. Enhanced surveys could identify which populations are most vulnerable to disturbance from human activities, and which should be highlighted for protection.

Deepcoral habitats of Glacier Bay National Park and Preserve, Alaska.

Glacier Bay National Park and Preserve (GBNPP) in Southeast Alaska is a system of glaciated fjords with a unique and recent history of deglaciation. As such, it can serve as a natural laboratory for studying patterns of distribution in marine communities with proximity to glacial influence. In order to examine the changes in fjord-based coral communities, underwater photo-quadrats were collected during multipurpose dives with a remotely operated vehicle (ROV) in March of 2016. Ten sites were chosen to represent the geochronological and oceanographic gradients present in GBNPP. Each site was surveyed vertically between 100 and 420 meters depth and photo-quadrats were extracted from the video strip transects for analysis. The ROV was equipped with onboard CTD which recorded environmental data (temperature and salinity), in order to confirm the uniformity of these characteristics at depth across the fjords. The percent cover and diversity of species were lowest near the glaciated heads of the fjords and highest in the Central Channel and at the mouths of the fjords. Diversity is highest where characteristics such as low sedimentation and increased tidal currents are predominant. The diverse communities at the mouths of the fjords and in the Central Channel were dominated by large colonies of the Red Tree Coral, Primnoa pacifica, as well as sponges, brachiopods, multiple species of cnidarians, echinoderms, molluscs and arthropods. The communities at the heads of the fjords were heavily dominated by pioneering species such as brachiopoda, hydrozoan turf, the encrusting stoloniferan coral Sarcodyction incrustans, and smaller colonies of P. pacifica. This research documents a gradient of species dominance from the Central Channel to the heads of the glaciated fjords, which is hypothesized to be driven by a combination of physical and biological factors such as glacial sedimentation, nutrient availability, larval dispersal, and competition.

Phenotypic Plasticity or a Reproductive Dead End? Primnoa pacifica (Cnidaria: Alcyonacea) in the Southeastern Alaska Region

Red tree corals (Primnoa pacifica) are abundant in the eastern Gulf of Alaska, from the glacial fjords of Southeast Alaska where they emerge to as shallow as 6 m, to the continental shelf edge and seamounts where they are more commonly found at depths greater than 150 m. This keystone species forms large thickets, creating habitat for many associated species, including economically valuable fishes and crabs, and so are important benthic suspension feeders in this region. Though the reproductive periodicity over 1.5 year period was reported in 2014 from a shallow fjord (Tracy Arm), this study examined reproductive ecologies from 8 sites – two within Glacier Bay National Park and Preserve, three on the continental shelf edge, one within Endicott Arm (Holkham Bay) and two time points from the Tracy Arm (Holkham Bay) study. Male reproductive traits were similar at all sites but there were distinct differences in oogenesis. Though fecundity mostly showed no significant difference between sites, there was a trend of increasing number of eggs with depth. Indeed, the average oocyte sizes from Tracy Arm (the shallowest site) was 105 m, whereas from Shutter Ridge (one of the deepest sites) the average size was 309 m. Moreover, the maximum oocyte size at Tracy Arm was 802 m, whereas it was 2120 m at Dixon Entrance, an almost 3-fold difference, and one not usually observed within a single species. We propose two theories to explain the observed differences, a) this species shows great phenotypic plasticity in reproductive ecology, adjusting to different environmental variables based on energetic need and potentially demonstrating micro-evolution; or b) the fjord sites are at a reproductive dead end, with the stress of shallow water conditions effectively preventing gametogenesis reaching full potential and likely limiting successful reproductive events from occurring, at least on a regular basis.

Eco-physiological responses of cold-water soft corals to anthropogenic sedimentation and particle shape

Anthropogenic disturbances in the marine environment, such as excessive sedimentation produced by mine tailing deposition, can affect the physiology and behavior of benthic fauna. Mine tailing particles are sharper than natural occurring sediments and could therefore cause more harmful effects. Cold-water coral ecosystems are among the habitats in danger of being affected by dumping of mine tailings. Cold-water corals have very slow growth rates but the habitats they provide support high levels of species richness and functional diversity. Two soft corals, cauliflower coral (Duva florida) and red tree coral (Primnoa resedaeformis), were chosen as model organisms to study the effects of excessive mine tailing sedimentation and the effect of particle shape in a three-month long experiment. The corals were exposed to a concentration of 8mgl−1 of two types of sediment, rough edged mine tailings (MT) and smooth edged spherical glass beads (GB). Glass beads were mimicking natural, smooth sediment, and both sediment types had a particle size distribution of 0–63μm. Sedimentation effects were investigated using 13C/12C isotope ratio to assess food intake, time-lapse images to determine the effects on tissue and behavior, and histological samples to identify and quantify particles inside the polyps. When exposed to MT, food intake decreased significantly in D. florida and increased significantly in P. resedaeformis. Duva florida exhibited a behavioral response under MT treatment, being contracted for prolonged periods. Primnoa resedaeformis lost a significant proportion of polyps under both treatments. Histology showed mine tailing particles of sizes <10μm embedded in the tissue of both species. The results suggest that sharp particles are more harmful than smooth edged particles to both species, in the size range studied. This should be considered when assessing the impacts of anthropogenic activities that increase sedimentation in benthic habitats.

Bayesian logistic mixed-effects modelling of transect data: relating red tree coral presence to habitat characteristics

Abstract The collection of continuous data on transects is a common practice in habitat and fishery stock assessments; however, the application of standard regression models that assume independence to serially correlated data is problematic. We show that generalized linear mixed models (GLMMs), i.e. generalized linear models for longitudinal data, that are normally used for studies performed over time can also be applied to other types of clustered or serially correlated data. We apply a specific GLMM for longitudinal data, a hierarchical Bayesian logistic mixed-effects model (BLMM), to a marine ecology dataset obtained from submersible video recordings of the seabed on transects at two sites in the Gulf of Alaska. The BLMM was effective in relating the presence of red tree corals (Primnoa pacifica; i.e. binary data) to habitat characteristics: the presence of red tree corals is highly associated with bedrock as the primary substrate (estimated odds ratio 9–19), high to very high seabed roughness (estimated odds ratio 3–5), and medium to high slope (estimated odds ratio 2–3). The covariate depth was less important at the sites. We also demonstrate and compare two methods of model checking: full and mixed posterior predictive assessments, the latter of which provided a more realistic assessment, and we calculate the variance partition coefficient for reporting the variation explained by multiple levels of the hierarchical model.

Assessing the ecological importance of red tree coral thickets in the eastern Gulf of Alaska

Abstract Red tree corals (Primnoa pacifica), the largest structure-forming gorgonians in the North Pacific Ocean, form dense thickets in some areas. These thickets are a dominant benthic habitat feature in the Gulf of Alaska (GOA), yet little is known about the ecosystems they support. In 2005, we used a submersible to study the ecology of thickets inside or near five small areas of the eastern GOA later designated in 2006 as habitat areas of particular concern (HAPCs)―areas closed to all bottom contact fishing. We show that red tree corals are keystone species in habitats where they form thickets (mean density 0.52 corals m−2)—the densest and largest thickets documented anywhere. Measured sponge densities (2.51 sponges m−2) were also among the highest documented anywhere. The corals and sponges in the study areas provide essential fish habitat for some fish species, and we show with logistic regression models modified with a scaled binomial variance that bedrock, while important habitat for some fish, is even more important when paired with corals and sponges. Red tree corals were not equally distributed with regard to habitat characteristics, and we show that their presence was correlated with bedrock substrate, moderate to high seabed roughness, and slope &amp;gt;10°. Most corals and sponges are vulnerable to disturbance from longlining, the principal bottom contact fishing in this region, but the larger corals and sponges are the most vulnerable. We observed evidence of infrequent recruitment events and a strong pulse of predation, apparently from fishing gear-induced trauma, that could exacerbate slow recovery of red tree corals from disturbance. Some red tree coral thickets are provided protection within designated HAPCs and some are not. Modifications to longline gear and an expanded network of HAPCs could help preserve these keystone species and the ecosystems they support.

Sexual reproduction and seasonality of the Alaskan red tree coral, Primnoa pacifica.

The red tree coral Primnoa pacifica is an important habitat forming octocoral in North Pacific waters. Given the prominence of this species in shelf and upper slope areas of the Gulf of Alaska where fishing disturbance can be high, it may be able to sustain healthy populations through adaptive reproductive processes. This study was designed to test this hypothesis, examining reproductive mode, seasonality and fecundity in both undamaged and simulated damaged colonies over the course of 16 months using a deepwater-emerged population in Tracy Arm Fjord. Females within the population developed asynchronously, though males showed trends of synchronicity, with production of immature spermatocysts heightened in December/January and maturation of gametes in the fall months. Periodicity of individuals varied from a single year reproductive event to some individuals taking more than the 16 months sampled to produce viable gametes. Multiple stages of gametes occurred in polyps of the same colony during most sampling periods. Mean oocyte size ranged from 50 to 200 µm in any season, and maximum oocyte size (802 µm) suggests a lecithotrophic larva. No brooding larvae were found during this study, though unfertilized oocytes were found adhered to the outside of polyps, where they are presumably fertilized. This species demonstrated size-dependent reproduction, with gametes first forming in colonies over 42-cm length, and steady oocyte sizes being achieved after reaching 80-cm in length. The average fecundity was 86 (±12) total oocytes per polyp, and 17 (±12) potential per polyp fecundity. Sub-lethal injury by removing 21–40% of colony tissue had no significant reproductive response in males or females over the course of this study, except for a corresponding loss in overall colony fecundity. The reproductive patterns and long gamete generation times observed in this study indicate that recruitment events are likely to be highly sporadic in this species increasing its vulnerability to anthropogenic disturbances.

Growth rate variation and potential paleoceanographic proxies in Primnoa pacifica: Insights from high-resolution trace element microanalysis

Red tree coral, Primnoa pacifica, is one of the more common habitat-forming deep-sea gorgonian corals in the northeast Pacific Ocean, growing in colonies up to 2m high and living for decades to hundreds of years. Growth characteristics of P. pacifica were studied in Dixon Entrance, northern British Columbia, and the Olympic Coast National Marine Sanctuary, Washington State, USA, based on samples collected in July 2008. To minimize the impact of scientific sampling on coral populations, only dead coral skeletons and dislodged live corals were collected. Ages and growth rates were measured using band counts, and checked against AMS-14C ages of gorgonin rings. Ba/Ca, Mg/Ca, Na/Ca and Sr/Ca ratios in the calcite cortex were measured using radial Secondary Ion Mass Spectrometer (SIMS) transects with a spot size of <20μm and separation distance of 25μm. Growth banding was consistent in width between the central mixed zone consisting of calcite and gorgonin and the dominantly calcite cortex. Average annual radial growth rate of the nine corals analysed ranged from 0.23 to 0.58mm/yr, with an average growth rate of 0.32mm/yr in Dixon Entrance and 0.36m/yr in OCNMS. These growth rates are slightly higher than P. pacifica growth rates from the Gulf of Alaska, and more than four times the growth rates of sister species Primnoa resedaeformis in the northwest Atlantic. Primary productivity is likely a more important driver of geographic variation in Primnoa growth rates than temperature or current strength. Both Dixon Entrance and OCNMS are areas with high primary productivity and strong tidal currents. Lack of post-Atomic Bomb radiocarbon in all but one of the gorgonin samples, and long radiocarbon reservoir ages in the Northeast Pacific, made radiocarbon-based verification of coral ages and growth rates difficult due to wide errors in calibrated age estimates. Mg/Ca and Sr/Ca ratios were inversely correlated in two of the three corals analyzed, and showed evidence of interannual variation. Mg/Ca ratios ranged from 70 to 136mmolmol−1, and Sr/Ca ratios from 2.041 to 3.14mmolmol−1. Previously published relationships between gorgonian calcite Mg/Ca and seawater temperature yielded average temperatures matching ambient measurements, but the intra- and inter-annual variation in apparent temperature based on the Mg/Ca ratios was more than double the observed variation in modern seawater temperature ranges in the region. Annual variation in Mg/Ca and Sr/Ca could be related to seasonal changes in precipitation efficiency, which is likely a function of short-term fluctuations in coral growth rate, in turn related to variation in primary productivity. Seasonal and interannual variations in food availability, driven by primary productivity, may affect skeletal growth rate, hence Mg/Ca and Sr/Ca ratios. Primnoid coral skeletal microgeochemistry probably records temporal changes in both temperature and primary productivity.

Mapping coral and sponge habitats on a shelf-depth environment using multibeam sonar and ROV video observations: Learmonth Bank, northern British Columbia, Canada

Efforts to locate and map deep-water coral and sponge habitats are essential for the effective management and conservation of these vulnerable marine ecosystems. Here we test the applicability of a simple multibeam sonar classification method developed for fjord environments to map the distribution of shelf-depth substrates and gorgonian coral- and sponge-dominated biotopes. The studied area is a shelf-depth feature Learmonth Bank, northern British Columbia, Canada and the method was applied aiming to map primarily non-reef forming coral and sponge biotopes. Aside from producing high-resolution maps (5m2 raster grid), biotope-substrate associations were also investigated. A multibeam sonar survey yielded bathymetry, acoustic backscatter strength and slope. From benthic video transects recorded by remotely operated vehicles (ROVs) six primary substrate types and twelve biotope categories were identified, defined by the primary sediment and dominant biological structure, respectively. Substrate and biotope maps were produced using a supervised classification mostly based on the inter-quartile range of the acoustic variables for each substrate type and biotope. Twenty-five percent of the video observations were randomly reserved for testing the classification accuracy. The dominant biotope-defining corals were red tree coral Primnoa pacifica and small styasterids, of which Stylaster parageus was common. Demosponges and hexactinellid sponges were frequently observed but no sponge reefs were observed. The substrate classification readily distinguished fine sediment, Sand and Bedrock from the other substrate types, but had greater difficulty distinguishing Bedrock from Boulders and Cobble. The biotope classification accurately identified Gardens (dense aggregations of sponges and corals) and Primnoa-dominated biotopes (67% accuracy), but most other biotopes had lower accuracies. There was a significant correspondence between Learmonth's biotopes and substrate types, with many biotopes strongly associated with only a single substrate type. This strong correspondence allowed substrate types to function as a surrogate for helping to map biotope distributions. Our results add new information on the distribution of corals and sponges at Learmonth Bank, which can be used to guide management at this location.

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.