Cerianthid anemones and burrowing megafauna in Atlantic mid bathyal mud
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| Researched by | Liam Matear & Laura Robson | Refereed by | This information is not refereed |
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Summary
UK and Ireland classification
Description
This biotope is characterized by burrowing anemones (Cerianthidae), unidentified hydroids (Hydrozoa) and unidentified tube worms (Sabellidae) on bioturbated mud with phytodetritus. Video observations suggest there are many large burrows supporting associated unknown megafauna, the rare occurrence of sea pens (Virgularia mirabilis) and also stalked sponges (Hyalonema) associated with this biotope.
Depth range
600-1300 mAdditional information
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Listed By
Sensitivity review
Sensitivity characteristics of the habitat and relevant characteristic species
Cerianthid anemones and burrowing megafauna occur in Atlantic mid bathyal mud and are characterized by Cerianthidae, Hydrozoa, Sabellidae, Hyalonema spp. and Virgularia mirabilis. The assessment is focused on Cerianthidae, Hydrozoa and Sabellidae, which are discussed in response to pressures individually. The loss or degradation of Cerianthidae, Hydrozoa, or, Sabellidae may result in direct changes to this biotope and therefore, the associated sensitivity values. Any differences between Cerianthidae, Hydrozoa and Sabellidae in response to pressures are highlighted, and the sensitivity of other biota is discussed where relevant.
The occurrence of Hyalonema spp. and Virgularia mirabilis is considered to be rare, therefore, these are not considered significant to the assessment of sensitivity. Further information on these species can be found in other biotope assessments on the MarLIN website.
Resilience and recovery rates of habitat
Cerianthid anemones are tube-forming, endobenthic cnidarians which inhabit marine reefs and benthic communities (Stampar et al., 2015). Individuals can occur on hard substrata, but predominantly inhabit soft, unconsolidated sediments (Stampar et al., 2015, 2014). Within the UK, Cerianthids have been recorded at varying depth ranges on both hard and soft sediment substrata, comprising bedrock, boulders, cobbles pebbles, muds and sands (Davies et al., 2014; Howell et al., 2010). Individuals are adapted to living in soft sediments in felted tubes made of cnidae, mucus and fragmented particles of sediment (Frey, 2012). Unlike other families within Ceriantharia, such as Arachnactidae, Cerianthidae are usually free standing, vertically orientated and unbranched in structure. Cerianthid tubes vary in thickness throughout life cycle stages and can reach up to 3 cm in adults or juveniles in later stages of development (Stampar et al., 2015). Tube development is initiated shortly after larvae settle on a suitable substratum, in which the ptychocyst, a tube forming cnida begins to form flat, overlapping fabric-like layers (Stampar et al., 2015). This style of tube development differs from that of species within Arachnactidae and Botrucnidiferidae, which utilise ptychocyst tubules to collect and trap passing sediment for tube formation (Stampar et al., 2015). In contrast to Actiniaria, Ceriantharia do not have longitudinal musculature, only circular, therefore, they use a combination of column constriction waves to displace unconsolidated material and ciliary movements of the column cells to enter the substratum (Stampar, S. personal communication). Ptychocyst filaments are released simultaneously by the polyp as it burrows, enabling for structural support to be given to the burrow walls (Stampar, S. pers. comm.). Burrows and tubes are considered permanent, in that the anemone does not voluntarily leave or move it (Bromley, 1996). However, if disturbed, organisms, including adults can leave and completely regenerate their tubes if necessary (Stampar et al., 2015). Should an organism become buried in sediment, tube formation ceases and the animal uses its tentacles to establish an escape trace, a new opening to the surface (Bromley, 1996). Organisms have been documented to create equilibrium structures, which develop as the individual maintains equilibrium with the seafloor throughout periods of sediment accretion (Bromley, 1996).
Cerianthids are microcarnivores that feed by using their marginal and inner tentacles to move prey to the pharynx for ingestion (Arai & Walder, 1973). Marginal tentacles can be activated through direct mechanical stimulation, or when labial tentacles are stimulated by water-soluble compounds (Arai & Walder, 1973). Furthermore, some organisms have been recorded to feed throughout night and daylight hours, without tidal rhythm, only withdrawing into their mucus tube if disturbed by external stimuli (Eleftheriou & Basford, 1983). Cerianthids have been documented to be both suspension and surface deposit feeders, although feeding may be inhibited and reduced under the force of high flow velocities, arising from strong swell conditions (Eleftheriou & Basford, 1983; Peters & Yevich, 1989). Feeding behaviour has been described as a series of continuous movements, searching for prey at wide angles, both above and in contact with surrounding sediment; the number of tentacles used is determined by the size of the prey (Eleftheriou & Basford, 1983).
Anthozoa are commonly either gonochoric or hermaphroditic. Mature gametes are shed into the coelenteron and are subsequently spawned into the water column through the mouth (Ruppert, 2004). Cerianthid zygotes develop into planula larvae, which begin metamorphosis once settled on a suitable substratum (Ruppert, 2004). Some evidence has documented larval development in Cerianthids, indicating that larvae are generally short-lived, with limited potential for dispersal. However, development time is genera or species specific, and certain Cerianthids can take up to several years to reach maturity (Hughes, 1998; Molodtsova, 2004). Furthermore, it has been highlighted previously that there is limited knowledge on a detailed understanding of sexual maturity and fecundity in Cerianthid species (Tyler-Walters & Hill, 2016).
Hydroids are predatory organisms, existing as either solitary polyps, or in large colonies joined by hydrocauli tubes, protected by a rigid exoskeleton called the perisarc (Moura et al., 2008). Hydrozoans vary in complexity and structure, comprising corneous, coriaceous and bilayered exoskeletons made from chitinous sheaths or calcareous coatings (Mendoza-Becerril et al., 2017). Most colonies are sessile, fixed to the seafloor by the stolon, a root-like structure formed from horizontal hydrocauli, which anchor the colony to the substratum (Moura et al., 2008). In colonial Hydroids, varying types of specialist polyps, or zooids, enable the individual to complete specific biological functions; for example, gastrozooids feed, dactylozoids predate, and gonozoids produce medusoids with gametes (Barnes, 1980). Hydroids are adapted to a diversity of ecological niches, inhabiting depths ranging from coastal to deep-water ecosystems and have been documented as widely distributed across UK and Irish waters, including deep-sea habitats in the UK South West Approaches (North East Atlantic) (Davies et al., 2014; “Hydrozoa,” n.d.; Moura et al., 2008). Organisms are capable of growing on both naturally occurring and artificial substrata, enabling conspicuous, passive dispersal through hydrochory and transport via anthropogenic substances through biofouling (Moura et al., 2008; Yan et al., 2006).
Hydroids are generally assumed to be carnivores, which predominantly feed on zooplankton using nematocysts on tentacles surrounding the mouth, or on the column of the hydranth (Gili & Hughes, 1995). Growth rates are dependent on food supply and are varied amongst Hydrozoans (Gili & Hughes, 1995). However, many are capable of exhibiting rapid growth due to the high volume of feeding hydranths, and therefore, high potential for predation (Gili & Hughes, 1995). Furthermore, temperature has been identified as a controlling factor capable of influencing the growth rate and stolon development, with increased growth exhibited as the temperature is increased (Berrill, 1949).
Hydroids exhibit rapid rates of recovery from disturbance through repair, asexual reproduction, and larval colonization. Sparks (1972) reviewed the regeneration abilities and rapid repair of injuries. Fragmentation of the hydroid provides a route for short distance dispersal, for example, each fragmented part of Sertularia cupressina can regenerate itself following damage (Berghahn & Offermann, 1999). New colonies of the same genotype may, therefore, arise from damage to existing colonies (Gili & Hughes, 1995). Many hydroid species also produce dormant, resting stages that are very resistant of environmental perturbation (Gili & Hughes 1995). Colonies can be removed or destroyed; however, the resting stages may survive attached to the substratum and provide a mechanism for rapid recovery (Kosevich & Marfenin, 1986; Cornelius, 1995a). The lifecycle of hydroids typically alternates between an attached solitary or colonial polyp generation and a free-swimming medusa generation. Planulae larvae produced by hydroids typically metamorphose within 24 hours and crawl only a short distance away from the parent plant (Sommer, 1992). Gametes liberated from the medusae (or vestigial sessile medusae) produce gametes that fuse to form zygotes and develop into free-swimming planula larvae (Hayward & Ryland, 1994) and are present in the water column between 2-20 days (Sommer, 1992). Generally, the planula larvae are short-lived and have been documented to take up to several days to settle (Moura et al., 2008). Rafting on floating debris as dormant stages or reproductive adults (or on ships hulls or in ship ballast water), together with their potentially long lifespan, may have allowed hydroids to disperse over a wide area in the long-term and explain the near cosmopolitan distributions of many hydroid species (Cornelius, 1992; Boero & Bouillon 1993). Hydroids are potential fouling organisms; rapidly colonizing a range of substrata placed in marine environments and are often the first organisms to colonize available space in settlement experiments (Gili & Hughes, 1995). For example, hydroids were reported to colonize an experimental artificial reef within less than 6 months, becoming abundant in the following year (Jensen et al., 1994). In similar studies, Obelia spp. recruited to the bases of reef slabs within three months and the slab surfaces within six months of the slabs being placed in the marine environment (Hatcher, 1998). Cornelius (1992) stated that Obelia spp. could form large colonies within a matter of weeks. In a study of the long-term effects of scallop dredging in the Irish Sea, Bradshaw et al. (2002) noted that hydroids increased in abundance, presumably because of their regeneration potential, good local recruitment and ability to colonize newly exposed substratum quickly. Cantero et al. (2002) describe fertility of Obelia dichotoma, Kirchenpaureria pinnata, Nemertesia ramosa in the Mediterranean as being year-round, whilst it should be noted that higher temperatures may play a factor in this year round fecundity. Bradshaw et al. (2002) observed that reproduction in Nemertesia antennina occurred regularly, with three generations per year. In addition, the presence of adults stimulated larval settlement, so that where adults remained, reproduction was likely to result in local recruitment.
Sabellidae, are a family of marine polychaete annelids, which can be identified by the prostomium and peristomium, which form a feathery tentacular crown, used for both respiration and feeding (Shumway et al., 1988; Tovar-Hernández & Salazar-Vallejo, 2006). Individuals are filter-feeding macroinvertebrates that feed using the branchial crown to collect suspended particulate matter, sorting particles by size for either ingestion or rejection (Bonar, 1972). Sabellids are tube-dwelling organisms on both hard and soft sediments such as sand, shell, cobbles and muds and are distributed across the UK and Ireland in both coastal and deep-sea habitats(Davies et al., 2014; McEuen et al., 1983; Nash & Keegan, 2003; “Sabellidae,” n.d.; Shumway et al., 1988).
Sabellidae have been documented to display a range of reproductive behaviours, including gamete dispersal, benthic egg mass deposition and brooding both outside and inside their tube (McEuen et al., 1983; Wilson, 1991). The gender of some species can be identified by dorsal aspect colouration, which is unique to males and females at varying life cycle stages (Nash & Keegan, 2003). Reproductive cycles have been documented for certain Sabellids, highlighting that spawning can be regulated by seasonal changes in water temperature, enabling species to produce larvae at specific times of the year (Nash & Keegan, 2003). For organisms which undergo external fertilization, synchronicity and coincidence of location between males and females is, therefore, integral to reproduction. Furthermore, in addition to temperature, the availability of food has been documented to regulate polychaete fecundity and reproductive mode, highlighting that impoverished organisms may have lower breeding rates than well-fed populations (Nash & Keegan, 2003; Tenore, 1977; Thorson, 1950). Larval development is initiated in egg cocoons, which have been documented in some Sabellids to hatch approximately eight days after deposition (McEuen et al., 1983). Larval stages are demersal and can take up to one week to settle before undergoing metamorphosis and tube development (McEuen et al., 1983). Tube formation is gradual and can result from burrowing, with sediment adhering to mucous sheets for tube development, or through the secretion of calcium carbonate (Glomerula only, non-UK genus) (Bonar, 1972; Perkins, 1991).
Resilience assessment. Hydroids are likely to recover from damage very quickly. Based on the available evidence, resilience for the hydroid species assessed is ‘High’ (recovery within two years) for any level of perturbation (where resistance is ‘None’, ‘Low’, ‘Medium’ or ‘High’). Sabellids, are probably able to recovery within a few years, although no direct evidence of recovery was found. However, the dominant characterizing species are the Cerianthids, and the ability of this biotope to recover will depend on the ability of Cerianthids to recover. However, there is very little information regarding the resilience of Cerianthids. Therefore, a resilience of ‘Medium’ (2 – 10 years) is suggested for all resistance levels (where resistance is ‘None’, ‘Low’, ‘Medium’) based on expert judgement, albeit at 'Low' confidence due to the lack of direct evidence for the characterizing species. An exception is made for permanent or ongoing (long-term) pressures where recovery is not possible as the pressure is irreversible, in which case resilience is assessed as ‘Very low’ by default.
Climate Change Pressures
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| Resistance | Resilience | Sensitivity | |
Global warming (extreme) [Show more]Global warming (extreme)Extreme emission scenario (by the end of this century 2081-2100) benchmark of:
EvidenceEvidence suggests that current deep-sea temperatures are generally low and stable (A. Levin & Le Bris, 2015; Pachauri et al., 2014). However, global mean sea surface temperatures have shown decadal increases of 0.11°C (1971-2010), and it has been reported by the Intergovernmental Panel on Climate Change (IPCC) that the effects of ocean warming are likely to impact depths ranging to 3000 m at the seafloor (A. Levin & Le Bris, 2015; Pachauri et al., 2014). Deep waters off the continental shelf (200 – 2,500 m) are predicted to see a lower temperature rise (ca 1°C) than shallow water habitats by the end of this century, regardless of the climate change scenario modeled (FAO (Fisheries and Aquaculture Organisation), 2019). Cerianthids can occur across wide temperature (8.36 – 11.51°C) and depth (238 – 1,070 m) ranges in UK deep-sea environments and Hydrozoans have been recorded in depths ranging to 8400 m, indicating adaptations to varied thermal niches throughout the water column (Davies et al., 2014; Stepanjants & Chernyshev, 2015). However, deep-sea environments are generally very stable and slow to change, therefore, a warming of 1°C may have deleterious impacts on sessile deep-sea organisms, resulting in changes in depth and latitudinal species distributions (Levin & Le Bris, 2015). Furthermore, studies have indicated that increased temperature can have pervasive stimulatory impacts on benthic invertebrate metabolism until lethal levels are reached (Byrne, 2011a). However, large knowledge gaps limit current understanding of the ecological feedbacks which may occur to Cerianthids as a result of climate change driven temperature increase. Elevated seawater temperature has been found to influence Hydrozoan metabolites; trigonelline abundance has been reduced in polyps exposed to warmer conditions, resulting in impaired polyp production (Boco et al., 2019). Therefore, restricted metabolic function resulting from increased temperature may reduce Hydrozoan abundances under projected ocean warming scenarios. In addition, Sabellid reproductive cycles have been documented to be directly influenced by temperature, with spawning periods and larval dispersal regulated by seasonal temperature variation (Nash & Keegan, 2003). Changes in reproduction periods have been documented for some Sabellids in relation to climate change, highlighting that warming conditions have altered settlement periods, and therefore, subsequent life cycle stages (Giangrande et al., 2010). However, due to the lack of data, the full impacts of seawater temperature increase as a result of climate change on Sabellids are poorly understood, and it is likely that changes to structure forming benthos will disrupt ecological equilibriums developed over long periods of time (Giangrande et al., 2010). Organisms that require temperature cues to synchronise external fertilization may therefore, experience inhibited reproduction as a result of temperature changes. Sensitivity Assessment. Under all scenarios (middle and high emission, and extreme scenarios), waters off the continental shelf are expected to increase marginally by approximately 1°C. Cerianthids have been recorded across broad temperature (8.36 – 11.51°C) and depth (238 – 1,070 m) ranges within UK deep-sea environments, suggesting that species within Cerianthidae can adapt to a diversity of thermal niches (Davies et al., 2014). However, sessile organisms, adapted to stable deep-sea environments may be at risk of pervasive stimulatory impacts arising from temperature increases of 1°C, resulting in changes in depth and latitudinal species distributions (Levin & Le Bris, 2015; Gibson et al., 2011). Furthermore, increased seawater temperature has been documented to inhibit metabolic function within some Hydrozoans, resulting in impaired polyp production, although evidence was not found documenting these impacts to species relevant to the UK deep-sea. Research has also indicated that temperature fluctuations resulting from climate change have altered reproduction and settlement periods for Sabellids, therefore synchronicity required for external fertilisation and reproduction may be impaired as a result of temperature change (Giangrande et al., 2010; Nash & Keegan, 2003). Data paucity was highlighted as a key limiting factor in understanding the ecological feedbacks which may arise from global warming (Byrne, 2011b). However, it is possible that temperature increases could impact all organisms considered characteristic of this biotope. Resistance for the biotope is assessed as ‘Medium’, but with 'Low' confidence. Global warming is considered a long-term, ongoing pressure, from which recovery is not possible, therefore, resilience is assessed as ‘Very low’ and overall sensitivity is assessed as ‘Medium’. | MediumHelp | Very LowHelp | MediumHelp |
Global warming (high) [Show more]Global warming (high)High emission scenario (by the end of this century 2081-2100) benchmark of:
EvidenceEvidence suggests that current deep-sea temperatures are generally low and stable (A. Levin & Le Bris, 2015; Pachauri et al., 2014). However, global mean sea surface temperatures have shown decadal increases of 0.11°C (1971-2010), and it has been reported by the Intergovernmental Panel on Climate Change (IPCC) that the effects of ocean warming are likely to impact depths ranging to 3000 m at the seafloor (A. Levin & Le Bris, 2015; Pachauri et al., 2014). Deep waters off the continental shelf (200 – 2,500 m) are predicted to see a lower temperature rise (ca 1°C) than shallow water habitats by the end of this century, regardless of the climate change scenario modeled (FAO (Fisheries and Aquaculture Organisation), 2019). Cerianthids can occur across wide temperature (8.36 – 11.51°C) and depth (238 – 1,070 m) ranges in UK deep-sea environments and Hydrozoans have been recorded in depths ranging to 8400 m, indicating adaptations to varied thermal niches throughout the water column (Davies et al., 2014; Stepanjants & Chernyshev, 2015). However, deep-sea environments are generally very stable and slow to change, therefore, a warming of 1°C may have deleterious impacts on sessile deep-sea organisms, resulting in changes in depth and latitudinal species distributions (Levin & Le Bris, 2015). Furthermore, studies have indicated that increased temperature can have pervasive stimulatory impacts on benthic invertebrate metabolism until lethal levels are reached (Byrne, 2011a). However, large knowledge gaps limit current understanding of the ecological feedbacks which may occur to Cerianthids as a result of climate change driven temperature increase. Elevated seawater temperature has been found to influence Hydrozoan metabolites; trigonelline abundance has been reduced in polyps exposed to warmer conditions, resulting in impaired polyp production (Boco et al., 2019). Therefore, restricted metabolic function resulting from increased temperature may reduce Hydrozoan abundances under projected ocean warming scenarios. In addition, Sabellid reproductive cycles have been documented to be directly influenced by temperature, with spawning periods and larval dispersal regulated by seasonal temperature variation (Nash & Keegan, 2003). Changes in reproduction periods have been documented for some Sabellids in relation to climate change, highlighting that warming conditions have altered settlement periods, and therefore, subsequent life cycle stages (Giangrande et al., 2010). However, due to the lack of data, the full impacts of seawater temperature increase as a result of climate change on Sabellids are poorly understood, and it is likely that changes to structure forming benthos will disrupt ecological equilibriums developed over long periods of time (Giangrande et al., 2010). Organisms that require temperature cues to synchronise external fertilization may therefore, experience inhibited reproduction as a result of temperature changes. Sensitivity Assessment. Under all scenarios (middle and high emission, and extreme scenarios), waters off the continental shelf are expected to increase marginally by approximately 1°C. Cerianthids have been recorded across broad temperature (8.36 – 11.51°C) and depth (238 – 1,070 m) ranges within UK deep-sea environments, suggesting that species within Cerianthidae can adapt to a diversity of thermal niches (Davies et al., 2014). However, sessile organisms, adapted to stable deep-sea environments may be at risk of pervasive stimulatory impacts arising from temperature increases of 1°C, resulting in changes in depth and latitudinal species distributions (Levin & Le Bris, 2015; Gibson et al., 2011). Furthermore, increased seawater temperature has been documented to inhibit metabolic function within some Hydrozoans, resulting in impaired polyp production, although evidence was not found documenting these impacts to species relevant to the UK deep-sea. Research has also indicated that temperature fluctuations resulting from climate change have altered reproduction and settlement periods for Sabellids, therefore synchronicity required for external fertilisation and reproduction may be impaired as a result of temperature change (Giangrande et al., 2010; Nash & Keegan, 2003). Data paucity was highlighted as a key limiting factor in understanding the ecological feedbacks which may arise from global warming (Byrne, 2011b). However, it is possible that temperature increases could impact all organisms considered characteristic of this biotope. Resistance for the biotope is assessed as ‘Medium’, but with 'Low' confidence. Global warming is considered a long-term, ongoing pressure, from which recovery is not possible, therefore, resilience is assessed as ‘Very low’ and overall sensitivity is assessed as ‘Medium’. | MediumHelp | Very LowHelp | MediumHelp |
Global warming (middle) [Show more]Global warming (middle)Middle emission scenario (by the end of this century 2081-2100) benchmark of:
EvidenceEvidence suggests that current deep-sea temperatures are generally low and stable (A. Levin & Le Bris, 2015; Pachauri et al., 2014). However, global mean sea surface temperatures have shown decadal increases of 0.11°C (1971-2010), and it has been reported by the Intergovernmental Panel on Climate Change (IPCC) that the effects of ocean warming are likely to impact depths ranging to 3000 m at the seafloor (A. Levin & Le Bris, 2015; Pachauri et al., 2014). Deep waters off the continental shelf (200 – 2,500 m) are predicted to see a lower temperature rise (ca 1°C) than shallow water habitats by the end of this century, regardless of the climate change scenario modeled (FAO (Fisheries and Aquaculture Organisation), 2019). Cerianthids can occur across wide temperature (8.36 – 11.51°C) and depth (238 – 1,070 m) ranges in UK deep-sea environments and Hydrozoans have been recorded in depths ranging to 8400 m, indicating adaptations to varied thermal niches throughout the water column (Davies et al., 2014; Stepanjants & Chernyshev, 2015). However, deep-sea environments are generally very stable and slow to change, therefore, a warming of 1°C may have deleterious impacts on sessile deep-sea organisms, resulting in changes in depth and latitudinal species distributions (Levin & Le Bris, 2015). Furthermore, studies have indicated that increased temperature can have pervasive stimulatory impacts on benthic invertebrate metabolism until lethal levels are reached (Byrne, 2011a). However, large knowledge gaps limit current understanding of the ecological feedbacks which may occur to Cerianthids as a result of climate change driven temperature increase. Elevated seawater temperature has been found to influence Hydrozoan metabolites; trigonelline abundance has been reduced in polyps exposed to warmer conditions, resulting in impaired polyp production (Boco et al., 2019). Therefore, restricted metabolic function resulting from increased temperature may reduce Hydrozoan abundances under projected ocean warming scenarios. In addition, Sabellid reproductive cycles have been documented to be directly influenced by temperature, with spawning periods and larval dispersal regulated by seasonal temperature variation (Nash & Keegan, 2003). Changes in reproduction periods have been documented for some Sabellids in relation to climate change, highlighting that warming conditions have altered settlement periods, and therefore, subsequent life cycle stages (Giangrande et al., 2010). However, due to the lack of data, the full impacts of seawater temperature increase as a result of climate change on Sabellids are poorly understood, and it is likely that changes to structure forming benthos will disrupt ecological equilibriums developed over long periods of time (Giangrande et al., 2010). Organisms that require temperature cues to synchronise external fertilization may therefore, experience inhibited reproduction as a result of temperature changes. Sensitivity Assessment. Under all scenarios (middle and high emission, and extreme scenarios), waters off the continental shelf are expected to increase marginally by approximately 1°C. Cerianthids have been recorded across broad temperature (8.36 – 11.51°C) and depth (238 – 1,070 m) ranges within UK deep-sea environments, suggesting that species within Cerianthidae can adapt to a diversity of thermal niches (Davies et al., 2014). However, sessile organisms, adapted to stable deep-sea environments may be at risk of pervasive stimulatory impacts arising from temperature increases of 1°C, resulting in changes in depth and latitudinal species distributions (Levin & Le Bris, 2015; Gibson et al., 2011). Furthermore, increased seawater temperature has been documented to inhibit metabolic function within some Hydrozoans, resulting in impaired polyp production, although evidence was not found documenting these impacts to species relevant to the UK deep-sea. Research has also indicated that temperature fluctuations resulting from climate change have altered reproduction and settlement periods for Sabellids, therefore synchronicity required for external fertilisation and reproduction may be impaired as a result of temperature change (Giangrande et al., 2010; Nash & Keegan, 2003). Data paucity was highlighted as a key limiting factor in understanding the ecological feedbacks which may arise from global warming (Byrne, 2011b). However, it is possible that temperature increases could impact all organisms considered characteristic of this biotope. Resistance for the biotope is assessed as ‘Medium’, but with 'Low' confidence. Global warming is considered a long-term, ongoing pressure, from which recovery is not possible, therefore, resilience is assessed as ‘Very low’ and overall sensitivity is assessed as ‘Medium’. | MediumHelp | Very LowHelp | MediumHelp |
Marine heatwaves (high) [Show more]Marine heatwaves (high)High emission scenario benchmark: A marine heatwave occurring every two years, with a mean duration of 120 days, and a maximum intensity of 3.5°C. Further detail. EvidenceMarine heatwaves caused by increased air-sea flux of heat are only expected to penetrate surface waters (≤ 50 m) (Cerrano et al., 2000, Garrabou et al., 2009; Dan Smale, pers. comms.). Cerianthid anemones and burrowing megafauna in Atlantic mid bathyal mud are deep-sea biotopes, relevant to the Atlantic mid bathyal zone, at depths of 600 – 1300 m. Therefore, the biotope will not be affected by changes arising from marine heatwaves, and the assessment at the pressure benchmark is ‘Not relevant’. | Not relevant (NR)Help | Not relevant (NR)Help | Not relevant (NR)Help |
Marine heatwaves (middle) [Show more]Marine heatwaves (middle)Middle emission scenario benchmark: A marine heatwave occurring every three years, with a mean duration of 80 days, with a maximum intensity of 2°C. Further detail. EvidenceMarine heatwaves caused by increased air-sea flux of heat are only expected to penetrate surface waters (≤ 50 m) (Cerrano et al., 2000, Garrabou et al., 2009; Dan Smale, pers. comms.). Cerianthid anemones and burrowing megafauna in Atlantic mid bathyal mud are deep-sea biotopes, relevant to the Atlantic mid bathyal zone, at depths of 600 – 1300 m. Therefore, the biotope will not be affected by changes arising from marine heatwaves, and the assessment at the pressure benchmark is ‘Not relevant’. | Not relevant (NR)Help | Not relevant (NR)Help | Not relevant (NR)Help |
Ocean acidification (high) [Show more]Ocean acidification (high)High emission scenario benchmark: a further decrease in pH of 0.35 (annual mean) and corresponding 120% increase in H+ ions , seasonal aragonite saturation of 20% of UK coastal waters and North Sea bottom waters, and the aragonite saturation horizon in the NE Atlantic, off the continental shelf, occurring at a depth of 400 m by the end of this century 2081-2100. Further detail EvidenceSustained anthropogenic emissions of carbon dioxide (CO2) into the atmosphere have contributed to a net flux of CO2 into the global ocean, resulting in pH decline through ocean acidification (Byrne, 2011b). However, despite the emerging evidence highlighting the variations in seawater pH due to climate change, understanding of the impacts of ocean acidification on deep-sea benthic ecology is limited (Danovaro, 2018). Decreased pH, and therefore shoaling of the calcium carbonate saturation horizon, may adversely impact UK sessile benthic communities (Danovaro et al., 2017). However, cerianthid anemones and sabellids (with the exception of Glomerula, a non-UK genus) are not calcifying organisms, and therefore, may not be impacted by inhibited calcification due to pH decline. In addition, evidence has indicated that some sabellids can maintain high homeostatic capacity and adaptability when exposed to elevated pCO2, although the species analysed in the studies were not UK, deep-sea organisms (Calosi et al., 2013; Del Pasqua et al., 2019). Furthermore, analyses focussing on taxa present in the North Atlantic have indicated that deep-sea communities containing cerianthids may not be adversely impacted by ocean acidification (Johnson et al., 2018). Knowledge gaps and data paucity were highlighted as key limitations in the study, therefore, conclusive assessments of pH decline impacts to such communities could not be made (Johnson et al., 2018). Ocean acidification has been documented to reduce calcification rates in some hydrocorals (de Barros Marangoni et al., 2017). In addition, hydrozoan metabolites required for cell protection from osmotic and thermal stress (betaine) have been found to be inhibited under extreme acidification (pH 7.7-7.75) as a result of climate change (Boco et al., 2019). However, such analyses have not been undertaken for deep-sea organisms within UK marine environments. In contrast, studies utilising over 40 years of Continuous Plankton Recorder (CPR) data from the North Sea have indicated a significant correlative relationship between increased Hydrozoan abundance and a temporal pH decline of 0.2 (Attrill et al., 2007; Fabry et al., 2008). Furthermore, some hydrozoans have been indicated to have high tolerances to wide pH variability, exhibiting complete mortality at pH 4 and 10, and with high survival rates (>50%) within the pH range of 5 – 8.5 (Gutierre, 2012). However, this assessment was not conducted for deep-sea species and tolerance ranges are likely to be species specific, therefore, it is possible that these findings cannot be inferred beyond the study. Sensitivity Assessment. Cerianthids and sabellids (with the exception of Glomerula, a non-UK genus) are not calcifying organisms, therefore, they are not considered to be impacted by the shoaling of the calcium carbonate saturation horizon due to ocean acidification (Danovaro et al., 2017). In addition, some sabellids have indicated the ability to adapt to acidification by maintaining high homeostatic capacity when exposed to seawater with elevated pCO2 concentrations (Calosi et al., 2013; Del Pasqua et al., 2019). Ocean acidification may limit calcification in some hydrocorals and has also been attributed to inhibiting metabolic function in hydrozoan metabolites required for cell protection against osmotic and thermal stress. (Boco et al., 2019; de Barros Marangoni et al., 2017). However, these impacts have not been documented in UK deep-sea environments and may be species-specific. Conversely, some hydrozoans have been found to be highly tolerant to wide pH ranges and CPR data have indicated a significant correlative relationship between increased hydrozoan abundance and increased acidification in the North Sea (Attrill et al., 2007; Fabry et al., 2008; Gutierre, 2012). However, data paucity has been identified as a key factor limiting understanding of acidification impacts to deep-sea biotopes (Johnson et al., 2018). Acidification at depth is driven by the solubility (thermohaline circulation) and biological pumps, which takes orders of magnitude longer than the air-to-surface-water CO2 exchange (Giering & Humphreys, 2017; Jones et al., 2014). For the middle (0.15 unit decrease in pH) and high-emission scenarios (0.35 unit decrease in pH), resistance is assessed as Medium as a precaution, based on the potential of decreases in pH limiting calcification in hydrocorals and inhibiting metabolic function. Ocean acidification is considered a long-term, ongoing pressure, from which recovery is not possible, therefore resilience is assessed as ‘Very low’ and overall sensitivity for this biotope is assessed as ‘Medium’. | MediumHelp | Very LowHelp | MediumHelp |
Ocean acidification (middle) [Show more]Ocean acidification (middle)Middle emission scenario benchmark: a further decrease in pH of 0.15 (annual mean) and corresponding 35% increase in H+ ions with no coastal aragonite undersaturation and the aragonite saturation horizon in the NE Atlantic, off the continental shelf, at a depth of 800 m by the end of this century 2081-2100. Further detail. EvidenceSustained anthropogenic emissions of carbon dioxide (CO2) into the atmosphere have contributed to a net flux of CO2 into the global ocean, resulting in pH decline through ocean acidification (Byrne, 2011b). However, despite the emerging evidence highlighting the variations in seawater pH due to climate change, understanding of the impacts of ocean acidification on deep-sea benthic ecology is limited (Danovaro, 2018). Decreased pH, and therefore shoaling of the calcium carbonate saturation horizon, may adversely impact UK sessile benthic communities (Danovaro et al., 2017). However, cerianthid anemones and sabellids (with the exception of Glomerula, a non-UK genus) are not calcifying organisms, and therefore, may not be impacted by inhibited calcification due to pH decline. In addition, evidence has indicated that some sabellids can maintain high homeostatic capacity and adaptability when exposed to elevated pCO2, although the species analysed in the studies were not UK, deep-sea organisms (Calosi et al., 2013; Del Pasqua et al., 2019). Furthermore, analyses focussing on taxa present in the North Atlantic have indicated that deep-sea communities containing cerianthids may not be adversely impacted by ocean acidification (Johnson et al., 2018). Knowledge gaps and data paucity were highlighted as key limitations in the study, therefore, conclusive assessments of pH decline impacts to such communities could not be made (Johnson et al., 2018). Ocean acidification has been documented to reduce calcification rates in some hydrocorals (de Barros Marangoni et al., 2017). In addition, hydrozoan metabolites required for cell protection from osmotic and thermal stress (betaine) have been found to be inhibited under extreme acidification (pH 7.7-7.75) as a result of climate change (Boco et al., 2019). However, such analyses have not been undertaken for deep-sea organisms within UK marine environments. In contrast, studies utilising over 40 years of Continuous Plankton Recorder (CPR) data from the North Sea have indicated a significant correlative relationship between increased Hydrozoan abundance and a temporal pH decline of 0.2 (Attrill et al., 2007; Fabry et al., 2008). Furthermore, some hydrozoans have been indicated to have high tolerances to wide pH variability, exhibiting complete mortality at pH 4 and 10, and with high survival rates (>50%) within the pH range of 5 – 8.5 (Gutierre, 2012). However, this assessment was not conducted for deep-sea species and tolerance ranges are likely to be species specific, therefore, it is possible that these findings cannot be inferred beyond the study. Sensitivity Assessment. Cerianthids and sabellids (with the exception of Glomerula, a non-UK genus) are not calcifying organisms, therefore, they are not considered to be impacted by the shoaling of the calcium carbonate saturation horizon due to ocean acidification (Danovaro et al., 2017). In addition, some sabellids have indicated the ability to adapt to acidification by maintaining high homeostatic capacity when exposed to seawater with elevated pCO2 concentrations (Calosi et al., 2013; Del Pasqua et al., 2019). Ocean acidification may limit calcification in some hydrocorals and has also been attributed to inhibiting metabolic function in hydrozoan metabolites required for cell protection against osmotic and thermal stress. (Boco et al., 2019; de Barros Marangoni et al., 2017). However, these impacts have not been documented in UK deep-sea environments and may be species-specific. Conversely, some hydrozoans have been found to be highly tolerant to wide pH ranges and CPR data have indicated a significant correlative relationship between increased hydrozoan abundance and increased acidification in the North Sea (Attrill et al., 2007; Fabry et al., 2008; Gutierre, 2012). However, data paucity has been identified as a key factor limiting understanding of acidification impacts to deep-sea biotopes (Johnson et al., 2018). Acidification at depth is driven by the solubility (thermohaline circulation) and biological pumps, which takes orders of magnitude longer than the air-to-surface-water CO2 exchange (Giering & Humphreys, 2017; Jones et al., 2014). For the middle (0.15 unit decrease in pH) and high-emission scenarios (0.35 unit decrease in pH), resistance is assessed as Medium as a precaution, based on the potential of decreases in pH limiting calcification in hydrocorals and inhibiting metabolic function. Ocean acidification is considered a long-term, ongoing pressure, from which recovery is not possible, therefore resilience is assessed as ‘Very low’ and overall sensitivity for this biotope is assessed as ‘Medium’. | MediumHelp | Very LowHelp | MediumHelp |
Sea level rise (extreme) [Show more]Sea level rise (extreme)Extreme scenario benchmark: a 107 cm rise in average UK by the end of this century (2018-2100). Further detail. EvidenceCerianthid anemones and burrowing megafauna in Atlantic mid bathyal mud are deep-sea biotopes, relevant to the Atlantic mid bathyal zone, at depths of 600 – 1300 m. Therefore, the biotope will not be affected by changes arising from sea-level rise, and the assessment at the pressure benchmark is ‘Not relevant’. | Not relevant (NR)Help | Not relevant (NR)Help | Not relevant (NR)Help |
Sea level rise (high) [Show more]Sea level rise (high)High emission scenario benchmark: a 70 cm rise in average UK by the end of this century (2018-2100). Further detail. EvidenceCerianthid anemones and burrowing megafauna in Atlantic mid bathyal mud are deep-sea biotopes, relevant to the Atlantic mid bathyal zone, at depths of 600 – 1300 m. Therefore, the biotope will not be affected by changes arising from sea-level rise, and the assessment at the pressure benchmark is ‘Not relevant’. | Not relevant (NR)Help | Not relevant (NR)Help | Not relevant (NR)Help |
Sea level rise (middle) [Show more]Sea level rise (middle)Middle emission scenario benchmark: a 50 cm rise in average UK sea-level rise by the end of this century (2081-2100). Further detail. EvidenceCerianthid anemones and burrowing megafauna in Atlantic mid bathyal mud are deep-sea biotopes, relevant to the Atlantic mid bathyal zone, at depths of 600 – 1300 m. Therefore, the biotope will not be affected by changes arising from sea-level rise, and the assessment at the pressure benchmark is ‘Not relevant’. | Not relevant (NR)Help | Not relevant (NR)Help | Not relevant (NR)Help |
Hydrological Pressures
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| Resistance | Resilience | Sensitivity | |
Temperature increase (local) [Show more]Temperature increase (local)Benchmark. A 5°C increase in temperature for one month, or 2°C for one year. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Temperature decrease (local) [Show more]Temperature decrease (local)Benchmark. A 5°C decrease in temperature for one month, or 2°C for one year. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Salinity increase (local) [Show more]Salinity increase (local)Benchmark. A increase in one MNCR salinity category above the usual range of the biotope or habitat. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Salinity decrease (local) [Show more]Salinity decrease (local)Benchmark. A decrease in one MNCR salinity category above the usual range of the biotope or habitat. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Water flow (tidal current) changes (local) [Show more]Water flow (tidal current) changes (local)Benchmark. A change in peak mean spring bed flow velocity of between 0.1 m/s to 0.2 m/s for more than one year. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Emergence regime changes [Show more]Emergence regime changesBenchmark. 1) A change in the time covered or not covered by the sea for a period of ≥1 year or 2) an increase in relative sea level or decrease in high water level for ≥1 year. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Wave exposure changes (local) [Show more]Wave exposure changes (local)Benchmark. A change in near shore significant wave height of >3% but <5% for more than one year. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Chemical Pressures
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| Resistance | Resilience | Sensitivity | |
Transition elements & organo-metal contamination [Show more]Transition elements & organo-metal contaminationBenchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Hydrocarbon & PAH contamination [Show more]Hydrocarbon & PAH contaminationBenchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Synthetic compound contamination [Show more]Synthetic compound contaminationBenchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Radionuclide contamination [Show more]Radionuclide contaminationBenchmark. An increase in 10µGy/h above background levels. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Introduction of other substances [Show more]Introduction of other substancesBenchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
De-oxygenation [Show more]De-oxygenationBenchmark. Exposure to dissolved oxygen concentration of less than or equal to 2 mg/l for one week (a change from WFD poor status to bad status). Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Nutrient enrichment [Show more]Nutrient enrichmentBenchmark. Compliance with WFD criteria for good status. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Organic enrichment [Show more] | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Physical Pressures
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| Resistance | Resilience | Sensitivity | |
Physical loss (to land or freshwater habitat) [Show more]Physical loss (to land or freshwater habitat)Benchmark. A permanent loss of existing saline habitat within the site. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Physical change (to another seabed type) [Show more]Physical change (to another seabed type)Benchmark. Permanent change from sedimentary or soft rock substrata to hard rock or artificial substrata or vice-versa. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Physical change (to another sediment type) [Show more]Physical change (to another sediment type)Benchmark. Permanent change in one Folk class (based on UK SeaMap simplified classification). Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Habitat structure changes - removal of substratum (extraction) [Show more]Habitat structure changes - removal of substratum (extraction)Benchmark. The extraction of substratum to 30 cm (where substratum includes sediments and soft rock but excludes hard bedrock). Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Abrasion / disturbance of the surface of the substratum or seabed [Show more]Abrasion / disturbance of the surface of the substratum or seabedBenchmark. Damage to surface features (e.g. species and physical structures within the habitat). Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Penetration or disturbance of the substratum subsurface [Show more]Penetration or disturbance of the substratum subsurfaceBenchmark. Damage to sub-surface features (e.g. species and physical structures within the habitat). Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Changes in suspended solids (water clarity) [Show more]Changes in suspended solids (water clarity)Benchmark. A change in one rank on the WFD (Water Framework Directive) scale e.g. from clear to intermediate for one year. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Smothering and siltation rate changes (light) [Show more]Smothering and siltation rate changes (light)Benchmark. ‘Light’ deposition of up to 5 cm of fine material added to the seabed in a single discrete event. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Smothering and siltation rate changes (heavy) [Show more]Smothering and siltation rate changes (heavy)Benchmark. ‘Heavy’ deposition of up to 30 cm of fine material added to the seabed in a single discrete event. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Litter [Show more]LitterBenchmark. The introduction of man-made objects able to cause physical harm (surface, water column, seafloor or strandline). Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Electromagnetic changes [Show more]Electromagnetic changesBenchmark. A local electric field of 1 V/m or a local magnetic field of 10 µT. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Underwater noise changes [Show more]Underwater noise changesBenchmark. MSFD indicator levels (SEL or peak SPL) exceeded for 20% of days in a calendar year. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Introduction of light or shading [Show more]Introduction of light or shadingBenchmark. A change in incident light via anthropogenic means. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Barrier to species movement [Show more]Barrier to species movementBenchmark. A permanent or temporary barrier to species movement over ≥50% of water body width or a 10% change in tidal excursion. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Death or injury by collision [Show more]Death or injury by collisionBenchmark. Injury or mortality from collisions of biota with both static or moving structures due to 0.1% of tidal volume on an average tide, passing through an artificial structure. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Visual disturbance [Show more]Visual disturbanceBenchmark. The daily duration of transient visual cues exceeds 10% of the period of site occupancy by the feature. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Biological Pressures
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| Resistance | Resilience | Sensitivity | |
Genetic modification & translocation of indigenous species [Show more]Genetic modification & translocation of indigenous speciesBenchmark. Translocation of indigenous species or the introduction of genetically modified or genetically different populations of indigenous species that may result in changes in the genetic structure of local populations, hybridization, or change in community structure. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Introduction or spread of invasive non-indigenous species [Show more]Introduction or spread of invasive non-indigenous speciesBenchmark. The introduction of one or more invasive non-indigenous species (INIS). Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Introduction of microbial pathogens [Show more]Introduction of microbial pathogensBenchmark. The introduction of relevant microbial pathogens or metazoan disease vectors to an area where they are currently not present (e.g. Martelia refringens and Bonamia, Avian influenza virus, viral Haemorrhagic Septicaemia virus). Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Removal of target species [Show more]Removal of target speciesBenchmark. Removal of species targeted by fishery, shellfishery or harvesting at a commercial or recreational scale. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Removal of non-target species [Show more]Removal of non-target speciesBenchmark. Removal of features or incidental non-targeted catch (by-catch) through targeted fishery, shellfishery or harvesting at a commercial or recreational scale. Further detail EvidenceNot assessed | Not Assessed (NA)Help | Not assessed (NA)Help | Not assessed (NA)Help |
Bibliography
Arai, M.N. & Walder, G.L., 1973. The feeding response of Pachycerianthus fimbriatus (ceriantharia). Comparative Biochemistry and Physiology Part A: Physiology, 44 (4), 1085-1092. DOI https://doi.org/10.1016/0300-9629(73)90246-6
Attrill, M.J., Wright, J. & Edwards, M., 2007. Climate-related increases in jellyfish frequency suggest a more gelatinous future for the North Sea. Limnology and Oceanography, 52 (1), 480-485. DOI https://doi.org/10.4319/lo.2007.52.1.0480
Barnes, R.D., 1980. Invertebrate Zoology, 4th ed. Philadelphia: Holt-Saunders International Editions.
Berrill, N.J., 1949. The polymorphic transformation of Obelia. Quarterly Journal of Microscopical Science, 90, 235-264.
Boco, S.R., Pitt, K.A. & Melvin, S.D., 2019. Extreme, but not moderate climate scenarios, impart sublethal effects on polyps of the Irukandji jellyfish, Carukia barnesi. Science of The Total Environment, 685, 471-479. DOI https://doi.org/10.1016/j.scitotenv.2019.05.451
- Bonar, D.B., 1972. Feeding and tube construction in chone mollis Bush (polychaeta, sabellidae). Journal of Experimental Marine Biology and Ecology, 9 (1), 1-18. DOI https://doi.org/10.1016/0022-0981(72)90002-0
Bromley, R.G., 2012. Trace Fossils: Biology, Taxonomy and Applications: Routledge.
- Byrne, M., 2011. Impact of ocean warming and ocean acidification on marine invertebrate life history stages: vulnerabilities and potential for persistence in a changing ocean. Oceanography and Marine Biology: An Annual Review, 49, 1-42.
Calosi, P., Rastrick, S.P.S., Lombardi, C., Guzman, H.J.d., Davidson, L., Jahnke, M., Giangrande, A., Hardege, J.D., Schulze, A., Spicer, J.I. & Gambi, M.-C., 2013. Adaptation and acclimatization to ocean acidification in marine ectotherms: an in situ transplant experiment with polychaetes at a shallow CO2 vent system. 368 (1627), 20120444. DOI https://doi.org/10.1098/rstb.2012.0444
Daly, M., Brugler, M.R., Cartwright, P., Collins, A.G., Dawson, M.N., Fautin, D.G., France, S.C., McFadden, C.S., Opresko, D.M., Rodriguez, E., Romano, S.L. & Stake, J.L., 2007. Phylogenetics of Hydroidolina (Hydrozoa: Cnidaria). Journal of the Marine Biological Association of the United Kingdom, 88 (8), 1663-1672.
Danovaro, R., 2018. Climate change impacts on the biota and on vulnerable habitats of the deep Mediterranean Sea. Rendiconti Lincei. Scienze Fisiche e Naturali, 29 (3), 525-541. DOI https://doi.org/10.1007/s12210-018-0725-4
Danovaro, R., Corinaldesi, C., Dell'Anno, A. & Snelgrove, P.V.R., 2017. The deep-sea under global change. Current biology: CB, 27 (11), R461-R465. DOI https://doi.org/10.1016/j.cub.2017.02.046
- Davies, J.S., Howell, K.L., Stewart, H.A., Guinan, J. & Golding, N., 2014. Defining biological assemblages (biotopes) of conservation interest in the submarine canyons of the South West Approaches (offshore United Kingdom) for use in marine habitat mapping. Deep Sea Research Part II: Topical Studies in Oceanography, 104, 208-229
De Barros Marangoni, L.F., Calderon, E.N., Marques, J.A., Duarte, G.A.S., Pereira, C.M., e Castro, C.B. & Bianchini, A.J.C.R., 2017. Effects of CO2-driven acidification of seawater on the calcification process in the calcareous hydrozoan Millepora alcicornis (Linnaeus, 1758). 36 (4), 1133-1141. DOI https://doi.org/10.1007/s00338-017-1605-6
Del Pasqua, M., Gambi, M.C., Caricato, R., Lionetto, M.G. & Giangrande, A., 2019. Effects of short-term and long-term exposure to ocean acidification on carbonic anhydrase activity and morphometric characteristics in the invasive polychaete Branchiomma boholense (Annelida: Sabellidae): A case-study from a CO2 vent system. Marine Environmental Research, 144, 203-212. DOI https://doi.org/10.1016/j.marenvres.2019.01.011
Eleftheriou, A. & Basford, D.J., 1983. The general behaviour and feeding of Cerianthus lloydii Gosse (Anthozoa, Coelenterata). Cahiers de Biologie Marine, 24, 147-158.
Fabry, V.J., Seibel, B.A., Feely, R.A. & Orr, J.C., 2008. Impacts of ocean acidification on marine fauna and ecosystem processes. ICES Journal of Marine Science, 65 (3), 414-432. DOI https://doi.org/10.1093/icesjms/fsn048
- Frey, R.W., 2012. The Study of Trace Fossils: A Synthesis of Principles, Problems, and Procedures in Ichnology: Springer Science & Business Media
Giangrande, A., Licciano, M., Musco, L. & Stabili, L., 2010. Shift in Sabella spallanzanii (Polychaeta, Sabellidae) spawning period in the Central Mediterranean Sea: a consequence of climate change? Mediterranean Marine Science, 11 (2), 373-380. DOI https://doi.org/10.12681/mms.86
- Giering, S. & Humphreys, M., 2017. Biological Pump. In White, W.M. (ed.) Encyclopedia of Earth Sciences Series: Springer International Publishing.
Gili, J-M. & Hughes, R.G., 1995. The ecology of marine benthic hydroids. Oceanography and Marine Biology: an Annual Review, 33, 351-426.
Gutierre, S.M.M., 2012. pH tolerance of the biofouling invasive hydrozoan Cordylophora caspia. Hydrobiologia, 679 (1), 91-95. DOI https://doi.org/10.1007/s10750-011-0855-5
Howell, K.L., Davies, J.S. & Narayanaswamy, B.E., 2010. Identifying deep-sea megafaunal epibenthic assemblages for use in habitat mapping and marine protected area network design. Journal of the Marine Biological Association of the United Kingdom, 90 (01), 33. DOI https://doi.org/10.1017/S0025315409991299
Hughes, D.J., 1998a. Sea pens & burrowing megafauna (volume III). An overview of dynamics and sensitivity characteristics for conservation management of marine SACs. Natura 2000 report prepared for Scottish Association of Marine Science (SAMS) for the UK Marine SACs Project., Scottish Association for Marine Science. (UK Marine SACs Project). Available from: http://ukmpa.marinebiodiversity.org/uk_sacs/pdfs/seapens.pdf
Johnson, D., Adelaide Ferreira, M. & Kenchington, E., 2018. Climate change is likely to severely limit the effectiveness of deep-sea ABMTs in the North Atlantic. Marine Policy, 87, 111-122. DOI https://doi.org/10.1016/j.marpol.2017.09.034
Jones, J., Duerdoth, C., Collins, A., Naden, P. & Sear, D., 2014. Interactions between diatoms and fine sediment. Hydrological Processes, 28 (3), 1226-1237.
Levin, L.A. & Le Bris, N., 2015. The deep ocean under climate change. Science, 350 (6262), 766-768. DOI https://doi.org/10.1126/science.aad0126
McEuen, F.S., Wu, B.L. & Chia, F.S., 1983. Reproduction and development of Sabella media, a sabellid polychaete with extratubular brooding. Marine Biology, 76 (3), 301-309. DOI https://doi.org/10.1007/BF00393033
Mendoza-Becerril, M.A., Marian, J.E.A.R., Migotto, A.E. & Marques, A.C., 2017. Exoskeletons of Bougainvilliidae and other Hydroidolina (Cnidaria, Hydrozoa): structure and composition. PeerJ, 5, e2964-e2964. DOI https://doi.org/10.7717/peerj.2964
Molodtsova, T.N., 2004. On the taxonomy and presumable evolutionary pathways of planktonic larvae of Ceriantharia (Anthozoa, Cnidaria). Hydrobiologia, 530/531, 261-266
Moura, C.J., Harris, D.J., Cunha, M.R. & Rogers, A.D., 2008. DNA barcoding reveals cryptic diversity in marine hydroids (Cnidaria, Hydrozoa) from coastal and deep-sea environments. Zoologica Scripta, 37 (1), 93-108. DOI https://doi.org/10.1111/j.1463-6409.2007.00312.x
Nash, R. & Keegan, B.F., 2003. Aspects of the feeding biology of the fanworm Bispira volutacornis (Polychaeta: Sabellidae). Journal of the Marine Biological Association of the United Kingdom, 83, 453-456.
- Pachauri, R.K., Allen, M.R., Barros, V.R., Broome, J., Cramer, W., Christ, R., Church, J.A., Clarke, L., Dahe, Q., Dasgupta, P., Dubash, N.K., Edenhofer, O., Elgizouli, I., Field, C.B., Forster, P., Friedlingstein, P., Fuglestvedt, J., Gomez-Echeverri, L., Hallegatte, S., Hegerl, G., Howden, M., Jiang, K., Jimenez Cisneroz, B., Kattsov, V., Lee, H., Mach, K.J., Marotzke, J., Mastrandrea, M.D., Meyer, L., Minx, J., Mulugetta, Y., O'Brien, K., Oppenheimer, M., Pereira, J.J., Pichs-Madruga, R., Plattner, G.K., Pörtner, H.-O., Power, S.B., Preston, B., Ravindranath, N.H., Reisinger, A., Riahi, K., Rusticucci, M., Scholes, R., Seyboth, K., Sokona, Y., Stavins, R., Stocker, T.F., Tschakert, P., van Vuuren, D. & van Ypserle, J.P., 2014. Climate Change 2014: Synthesis Report. Contribution of Working Groups I, II and III to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change, Geneva, Switzerland: IPCC. Available from: https://www.ipcc.ch/report/ar5/syr/
- Peters, E.C. & Yevich, P.P., 1989. Histopathology of Ceriantheopsis americanus (Cnidaria: Ceriantharia) exposed to Black Rock. Diseases of aquatic organisms, 7, 137-148
Ruppert, E.E. & Barnes, R.D., 1994. Invertebrate zoology (6th ed.). Fort Worth, USA: Saunders College Publishing.
- Shumway, S.E., Bogdanowicz, C. & Dean, D., 1988. Oxygen consumption and feeding rates of the sabellid polychaete, Myxicola infundibulum (Renier). Comparative Biochemistry and Physiology Part A: Physiology, 90 (3), 425-428
Stampar, S.N., Beneti, J.S., Acuña, F.H. & Morandini, A.C., 2015. Ultrastructure and tube formation in Ceriantharia (Cnidaria, Anthozoa). Zoologischer Anzeiger - A Journal of Comparative Zoology, 254, 67-71. DOI https://doi.org/10.1016/j.jcz.2014.11.004
Stepanjants, S.D. & Chernyshev, A.V., 2015. Deep-sea epibiotic hydroids from the abyssal plain adjacent to the Kuril–Kamchatka Trench with description of Garveia belyaevi sp. nov. (Hydrozoa, Bougainvilliidae). Deep Sea Research Part II: Topical Studies in Oceanography, 111, 44-48. DOI https://doi.org/10.1016/j.dsr2.2014.07.015
Tenore, K.R., 1977. Growth of Capitella capitata cultured on various levels of detritus derived from different sources. Limnology and Oceanography, 22 (5), 936-941.
Thorson, G., 1950. Reproductive and larval ecology of marine bottom invertebrates. Biological Reviews, 25, 1-45.
- Tovar-Hernández, M.A. & Salazar-Vallejo, S., 2006. Sabellids (Polychaeta: Sabellidae) from the Grand Caribbean. Zoological studies, 45, 24-66
- Tyler-Walters, H. & Hill, J.M., 2016. Philine quadripartita and Virgularia mirabilis in soft stable infralittoral mud. In Tyler-Walters, H. and Hiscock, K. (eds.). Marine Life Information Network: Biology and Sensitivity Key Information Reviews (online): Marine Biological Association of the UK, Plymouth
- Wilson, W.H., 1991. Sexual reproductive modes in Polychaetes: Classification and diversity. Bulletin of Marine Science, 48 (2), 500-516
Yan, T., Yan, W., Dong, Y., Wang, H., Yan, Y. & Liang, G., 2006. Marine fouling of offshore installations in the northern Beibu Gulf of China. International Biodeterioration & Biodegradation, 58, 99-105. DOI https://doi.org/10.1016/j.ibiod.2006.07.007
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Last Updated: 21/11/2019
