Geodia and other massive sponges on Atlanto-Arctic upper bathyal mixed sediment

Summary

UK and Ireland classification

Description

This biotope has been found at approximately 500 m on the eastern slopes of the Faroe-Shetland Channel.  It consists principally of rather small sponge specimens but that practically carpet the seafloor. Large sponges (tens of centimetres in diameter) are, however, common. The area where the community is observed experiences temperature fluctuations between <0°C to >8°C and heightened current speeds as a result of internal tides between Arctic and Atlantic environmental data. This assemblage is also known as Boreal 'Ostur'. The same assemblage was recorded on sand substratum, but associated infaunal species are likely to differ. Characterizing species listed refer to all Geodia and other massive sponge assemblages not just those found associated with the zone and substratum  specified in this biotope. (Information from JNCC, 2015; Parry et al., 2015).

Depth range

300-600 m

Additional information

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Sensitivity reviewHow is sensitivity assessed?

Sensitivity characteristics of the habitat and relevant characteristic species

The characterizing species for assemblages on both coarse and mixed sediments are Geodia spp., which could include Geodia barretti, Geodia macandrewii, Geodia atlantica and Geodia phlegraei, and the massive demosponges Stryphnus ponderosus and Stelletta normani. Loss of these predominant species making up the biotopes would result in loss or degradation of the biotope. Therefore, the sensitivities of the biotopes are dependent on the sensitivities of these species. Evidence for the specific Geodia species listed above has been used for the sensitivity assessment where possible. However, these are all suitable proxy species for each other and the overall sensitivity of these biotopes has been heavily based on the deep-sea Geodia genus as a whole.  In addition, where evidence of the effect of pressures on other demosponges deep-sea species, such as Radiella or Polymastia, is available, these species have been used in the assessment, and this is reflected in the confidence score.

Resilience and recovery rates of habitat

Geodia barretti, Geodia macandrewii, Geodia atlantica and Geodia phlegraei, of the class Demospongiae (sub-order Astrophorina) form aggregations on the outer shelf and upper slope over large areas of the NE Atlantic (Spetland et al., 2007). The sub-order Astrophorina, which also includes Stryphnus ponderosus and Stelletta normani, have a degree of structural flexibility, due to a non-rigid structural matrix of spicules and spongin fibres (mesohyl) (Uriz et al., 2003). Geodia barretti can be found on both hard and sedimentary substrata (Henry & Roberts, 2014; Klitgaard & Tendal, 2004). In high-energy environments, Geodia barretti is found attached to stable substrata, while in areas with lower energy it is not attached, but incorporates small stones in order to maintain negative buoyancy (Tjensvoll et al., 2013). There are numerous factors that control the spatial distribution of sponges, including substratum, food availability, exposure, competition, predation, temperature and suitability for larvae (Kutti et al., 2013).

Deep-sea demosponge grounds dominated by the families Geodiidae and Ancordinidae (Stryphnus ponderosus and Stelletta normani) are known as boreal ‘ostur’, and occur in the Faroes, Arctic and all over the north-east Atlantic (north of 60°N) (Klitgaard & Tendal, 2004). The distribution of deep-sea sponge aggregations has historically been inferred from fisheries by-catch and anecdotal knowledge (Klitgaard & Tendal, 2004). Scientific effort has gone into locating, describing and protecting these habitats only recently (Beazley et al., 2013; McIntyre et al., 2016; Roberts et al., 2018). In addition, several studies have modelled the distribution of deep-sea sponge assemblages, providing useful insight into potential factors that control the suitability of habitat for deep-sea sponges (Howell et al., 2016; Knudby et al., 2013). The OSPAR definition of deep-sea sponge aggregations identifies the 250 m water depth contour as the upper limit of aggregations (Christiansen, 2010), however boreal ‘ostur’ have also been recorded off the Norwegian south coast at depths of between 100-300 m (Klitgaard & Tendal, 2004).

Geodia and other massive sponges can form dense assemblages of up to 0.4 m2, where they dominate the benthic macrofauna in body size, abundance and, often, total invertebrate biomass (Klitgaard & Tendal, 2004; Kutti et al., 2013; Roberts et al., 2018). Surveys of these grounds in Faroese waters have found some individual specimens of Geodia barretti, Geodia macandrewii, Geodia atlantica and Stryphnus ponderosus exceeding 70 cm in diameter and 24 kg wet weight (Klitgaard & Tendal, 2004). Leys et al. (2018) recorded one specimen of Geodia barretti with a wet weight of 36.7 kg, and Kutti et al. (2013) found specimens of Geodia atlantica that were 128cm in width.

Of the characterizing species for this biotope, specific reproductive evidence is only available for Geodia barretti, which is oviparous dioecious and an annual spawner, with one or two periods of gamete release each year (Spetland et al., 2007). Individuals are thought to reproduce simultaneously within the same local population, within a restricted time period (Spetland et al., 2007). Asexual reproduction is not thought to be possible in Geodia barretti and it is noted that hermaphroditism is very rare in Astrophoridian sponges (Scalera Liaci & Sciscioli, 1970). In two Scandinavian fjords, the onset of reproduction coincides with the phytoplankton blooms and the release of gametes is likely to occur when sedimentation of particulate organic matter is at its highest after the spring phytoplankton blooms, i.e. in early summer, but also in October after the autumn phytoplankton blooms (Spetland et al., 2007). Specific information for other Geodia species, Stryphnus ponderosus or Stelletta normani was not available. Geodia spp. larvae, like the majority of other sponge larvae, are non-feeding and short-lived (remaining in the water column for only a few hours) (Maldonado & Bergquist, 2002, cited in Knudby et al., 2013), and they settle in the vicinity of parental populations (Mariani et al., 2003, cited in Knudby et al., 2013). Knudby et al. (2013) have shown that there is low connectivity between different areas of Geodia spp. (>1500 km apart), resulting from short-range larval dispersal and high larval retention. It is, therefore, possible that populations are highly inbred and potentially locally adapted (Knudby et al., 2013).

There is limited information on the longevity and age of sexual maturity in deep-sea sponges. The capture of individuals that are in transition from immaturity for ageing is exceptionally difficult. Studies of the growth rates of deep-sea sponges are very rare due to the difficulty of obtaining direct time-series observations (Serpetti et al., 2014). There is strong evidence of extreme lifespans in deep-sea glass sponges (Hexactinellids), ranging from 220 years (Leys & Lauzon, 1998) to 440 years (Fallon et al., 2010). It is possible that the characterizing sponge species also have long–lifespans, but there is no direct evidence on lifespan in the deep-sea Geodia spp., Stryphnus ponderosus or Stelletta normani.

Hoffmann et al. (2003) reported that there was no measurable change in the size or shape of a specimen of Geodia barretti over a two-year period in a Norwegian fjord (personal observation). Klitgaard & Tendal (2004) further suggested that the dominant species in boreal ‘ostur’ communities (i.e. Geodia spp.) are likely to be slow-growing, taking at least several decades to reach their large sizes. Deep-sea Hexactinellid sponges, for example, have been calculated to have a linear extension rate of around 2.9 mm/yr for some species (Fallon et al., 2010), whilst others have an average growth rate of 1.98 cm/yr (Leys & Lauzon, 1998). In the characterizing species Geodia barretti, Hoffmann et al. (2003) found that fragments on the inner lining (choanosomal) were able to regenerate body tissue (cortex), and after one year of cultivation in the field, the sponges had increased in weight by 40%. Kutti et al. (2015) also cultivated Geodia barretti using open sea aquaculture in the field, followed by two-months of adaptation in a laboratory (the same method as Hoffmann et al., 2003). Beginning from sponge tissue of 4 x 4 x 4 cm in size, within eight months of cultivation the sponge explants were pumping and had respiration rates similar to that of full-grown individuals, showing cortex regeneration had taken place (Kutti et al., 2015). Survival rate during cultivation in the field was 50% (Kutti et al., 2015).

Freese (2001) studied the recovery of deep-sea sponges off Alaska, one year after experimental trawls. This included sponges initially thought to be Geodia sp. but later re-identified as likely Poecillastra tenuilaminaris, of the same sub-order Astrophorina (Malecha & Heifetz, 2017). Analysis of video data from three of the re-visited trawl transects showed a mean percentage of 56.3% of Geodia sp. lying on the bottom either torn or intact, 40.6% undamaged, and 3.1% upright but torn (Freese, 2001). None of the damaged sponges displayed signs of regrowth or recovery a year after the trawl event but most of the knocked over sponges or pieces of sponge that had been torn off still appeared viable (Freese, 2001). In a follow-up study, 13-years post-trawling, Malecha & Heifetz (2017) found long-term damage and potential delayed mortality to sponges with an average of 31.7% lower density of all sponge species in trawled vs. reference transects. An average of 57.1% of Poecillastra tenuilaminaris (re-identified from Geodia sp.) were damaged (either lying on the bottom or torn) in trawled areas, vs. 16.7% in reference areas.

The feeding method of the characterizing species, in common with many sponges, is through filter feeding. Geodia barretti has been shown to have a very low specific filtration and low respiration rates in comparison to other sponges (Leys et al., 2018). Other cold-water sponges share similar low specific respiration rates, and these lower rates are thought to be dictated by water temperature and habitat (Leys et al., 2018). Sponges are known to be important ecosystem engineers (Bell, 2008), contributing to biogeochemical processes and, in the deep sea, are important in carbon cycling. One study suggests that within an area of 300 km2, a population of Geodia barretti with an average biomass of 1.4 kg WW m-2 could filter 250 million mof water and consume 60 t of carbon every 24 hours (Kutti et al., 2013). Clearly, this significant volume of water filtration will have implications for the biotope in the event of changing sponge abundances or pumping rates. In addition, stress effects are known to affect filter-feeding in deep-sea sponges (Robertson et al., 2017).

In addition, the taxonomy of the characterizing species is somewhat fluid, such as the division of Geodia phlegraei into Geodia phlegraei and Geodia parva (Cárdenas et al., 2013).  However, the functionality of these species is likely to be the same and, therefore, species-specific taxonomy probably has no implications for the sensitivity assessment.     

Resilience assessment.  Where resistance is ‘None’ or ‘Low’, and an element of habitat recovery is required, resilience is assessed as ‘Very low’ (>25 years). This is based upon the slow growth rates of Geodia sp. (i.e., several decades; Klitgaard and Tendal, 2004), likely extreme lifespans (as in other deep-sea sponges; Fallon et al., 2010; Leys and Lauzon, 1998), low connectivity (i.e., short-range larval dispersal and high larval retention; Knudby et al., 2013), and their low respiration and filtration rates (Leys et al., 2018). Furthermore, studies have shown that recovery from sedimentation events was negligible over a 10-year period (Jones et al., 2012) with significantly delayed mortality and stress effects still evident in the deep-sea sponge communities after 13-years following trawling impact (Malecha & Heifetz, 2017). It is also important to note the lack of basic information on Geodia sp., and not knowing the population structure (i.e., how larvae and juveniles will be affected). The confidences associated with these scores are ‘High’ for Quality of Evidence, ‘Medium’ for Applicability of Evidence (studies are inferred from species knowledge from other ecosystems, e.g. Alaska) and ‘High’ for Degree of Concordance.

Where resistance of the characterizing species is ‘Low’ or ‘Medium’, and the habitat has not been altered, resilience is assessed as ‘Low’ (10-25 years). Despite Geodia barretti exhibiting annual spawning (Spetland et al., 2007), with surviving sponges potentially providing a nucleus for population recovery (Serpetti et al., 2014), the community will take longer than 10 years to recover to the pre-impact status due to the slow-growing rates of Geodia sp. (i.e., several decades; Klitgaard and Tendal, 2004). Furthermore, the likely extreme lifespans (as in other deep-sea sponges; Fallon et al., 2010; Leys and Lauzon, 1998), low connectivity (i.e., short-range larval dispersal and high larval retention; Knudby et al., 2013), and their low respiration and filtration rates (Leys et al., 2018) will further slow their recovery rates.  Experiments have shown that Geodia barretti explants exhibit pumping and respiration rates similar to full-grown individuals after eight months of cultivation in the field (Kutti et al., 2015; Hoffmann et al., 2003), however, recovery to their former structural complexity will likely take much longer due to their slow growth rates (i.e., several decades; Klitgaard & Tendal, 2004). It is important to note the lack of basic information on Geodia sp., or its population structure (i.e., how larvae and juveniles will be affected). The confidences associated with these scores are ‘High’ for quality of evidence, ‘Medium’ for applicability of evidence (studies are from Norwegian fjords and north-west Atlantic) and ‘High’ for degree of concordance.

Hydrological Pressures

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ResistanceResilienceSensitivity
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

Evidence

Modelling studies have shown temperature to be an important driver behind deep-sea sponge distribution (Howell et al., 2016; Knudby et al., 2013), while direct measurements have shown stable, low bottom temperatures in Arctic sponge communities (Roberts et al., 2018).  The area of the Faroe-Shetland Channel where the biotope is observed in the UK experiences natural fluctuations in temperature from <0°C to >8°C (Parry et al., 2015). The characterizing species Geodia barretti normally lives in deep-sea waters where temperatures are 8-9°C (Leys et al., 2018), however, Henry & Roberts (2014) mention that boreal ‘ostur’ occurs in sub-zero temperatures on the northern side of the Wyville-Thomson Ridge.

Geodia barretti also occurs in the Barents Sea where bottom temperatures average 4.8°C (Kędra et al., 2017).  Specimens were found in the Mediterranean Sea, at a depth of 167 m, where the water temperature was around 13°C (Cárdenas et al., 2013). In the Faroe-Shetland Channel, Kazanidis et al. (2019) observed that the transects with the highest densities of sponges had temperatures that ranged from 6.52 to 8.98°C. Kazanidis et al. (2019) also found that temperature was a significant variable in explaining variations in sponge density.

There have been concerns that real-world temperature increases can be deleterious.  Mass mortality of Geodia barretti in 2006 and 2008 at the Tisler reef, Norway was concurrent with ~4°C bottom temperature increases over 24 hr periods, for up to two weeks (Guihen et al., 2012). However, in vitro experiments were carried out to replicate this event and no visible signs of stress were found (Strand et al., 2017). Although increased respiration rate was noted, the sponge’s microbiome remained stable, no mortality was observed and respiration rates returned to normal on return to control temperatures (Strand et al., 2017). This could suggest that Geodia barretti is resistant to a temperature increase at benchmark level and that other factors must have been complicit in the mass mortality seen at the Tisler reef (Strand et al., 2017). However, the confidence in the degree of concordance for this assessment has been recorded as ‘Low’ to reflect this uncertainty.  

The study by Strand et al. (2017) is not alone in suggesting that temperature, along with other stressors, may create an additive effect that may be more deleterious to the characterizing species. Scanes et al. (2018) found that the multiple stressors of increased temperature (+5°C) and suspended sediment (simulated inert mine tailings, at 10 mg/l) on Geodia atlantica interacted to cause an increase in the release of nitrogen, indicating a higher energy demand. The temperature increase itself also caused increased respiration and a greater percentage of unstable lysosomes (reduced cellular health), which may also be a result of an increase in energetic demand, as well as stress (Scanes et al., 2018). Furthermore, an increase in silicate uptake (used for biomineralization to produce their skeletons) was observed with elevated temperature (13°C) after 33 days (Scanes et al., 2018). However, this is opposite to previous and subsequent observations.  For example, Strand et al. (2017) found no effect on silicate uptake with warming.  The result of the multiple stressors (warming and suspended sediments) must be interpreted carefully. The authors found that there were complex interactive effects on the species under study. Unpicking these relationships to evaluate temperature change effects is not straightforward.

Overall, if the biotopes occur in areas corresponding to the lower limit or middle of their temperature range, then they are probably able to tolerate a long-term increase in temperature of 2°C. However, if the biotopes occur at the upper limit of their temperature range (i.e. 8.98°C; Kazanidis et al., 2019), then they probably cannot tolerate an increase in temperature of 2°C. Although specimens of Geodia barretti have been found in the Mediterranean at temperatures of 13°C (Cárdenas et al., 2013), these records do not represent the biotopes of interest (i.e. Geodia and other massive sponges on Atlanto-Arctic upper bathyal coarse or mixed sediment), so it cannot be assumed that the biotopes would survive at these temperatures. It is likely that only biotopes at the lowest limit of their temperature range will tolerate a short-term increase in temperature of 5°C, and the above studies have shown potentially deleterious effects caused by 4 or 5°C increases in temperature. It is important to note that an increase in temperature may also cause a decrease in water oxygen concentration, however, this pressure is addressed separately (see the ‘De-oxygenation’ pressure).

Sensitivity assessment. The biotopes are associated with a range of bottom water temperatures, however increasing temperature may have an effect on Geodia and other sponge species making up the biotopes, particularly when in combination with other stressors. Therefore, resistance is assessed as ‘Medium’, resilience as ‘Low’ and the sensitivity is assessed as ‘Medium’.

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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

Evidence

There is a range of evidence surrounding the effect of temperature change on the characterizing species and on sponge aggregations in other regions. Modelling studies have shown temperature to be an important driver behind deep-sea sponge distribution (Howell et al., 2016; Knudby et al., 2013) while direct measurements have shown stable, low bottom temperatures in Arctic sponge communities (Roberts et al., 2018). The area of the Faroe-Shetland Channel where the biotope is observed in the UK experiences natural fluctuations in temperature from <0°C to >8°C (Parry et al., 2015). The characterizing species Geodia barretti normally lives in deep-sea waters where temperatures are 8-9°C (Leys et al., 2018), however, Henry & Roberts (2014) also mention that boreal ‘ostur’ occurs in sub-zero temperatures on the northern side of the Wyville-Thomson Ridge. Geodia barretti also occurs in the Barents Sea where bottom temperatures average 4.8°C (Kędra et al., 2017). Specimens were found in the Mediterranean Sea, at a depth of 167 m, where the water temperature was around 13°C (Cárdenas et al., 2013). In the Faroe-Shetland Channel, Kazanidis et al. (2019) observed that the transects with the highest densities of sponges had temperatures that ranged from 6.52 to 8.98°C. Kazanidis et al. (2019) also found that temperature was a significant variable in explaining variations in sponge density.

Direct laboratory experiments on deep-sea representatives of the genera Polymastia and Radiella found that a decrease from 6°C to 0 and 3°C had no significant effect on feeding rates, although there was a decrease in filtration rates for Polymastia but not at a significant level (Robertson et al., 2017). They concluded that naturally occurring temperature shifts (such as those caused by the Labrador current in Newfoundland where their study was located) were well tolerated by sponges (Robertson et al., 2017). The study by Robertson et al. (2017) supports earlier conclusions that sponge communities could tolerate changes in thermal regime better than other groups, such as cold-water corals during El Niño Southern Oscillation (ENSO) events  (Kelmo et al., 2013).

Overall, if the biotopes occur in an area corresponding to the upper limit or middle of their temperature range, then they are probably able to tolerate a long-term decrease in temperature of 2°C. However, if the biotopes occur at the lower limit of their temperature range, then they probably cannot tolerate a decrease in temperature of 2°C. At the upper limit of their temperature range (8.98°C; Kazanidis et al., 2019) the biotopes are likely to tolerate a short-term decrease in temperature of 5°C, as this falls within the know temperature range of the biotopes.

Sensitivity assessment: Geodia barretti occurs at a range of temperatures in deep-sea waters from sub-zero to around 8.98°C (Henry & Roberts, 2014; Kazanidis et al., 2019) and direct laboratory experiments on other deep-sea species have shown that a decrease in temperature at the benchmark level would not have a significant effect. Therefore, resistance is assessed as ‘High’, resilience as ‘High’, and the biotope is assessed as ‘Not sensitive’ at the benchmark level.

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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

Evidence

Changes in salinity are unlikely due to the depth at which the biotopes are found, combined with the distance from shore and the low potential for brine or freshwater discharge. However, salinity (along with other factors) is an important controller of modelled deep-sea sponge distribution (Howell et al., 2016; Knudby et al., 2013). Many studies examining the distribution of deep-sea sponges are limited to recording the conditions present at natural sponge aggregations. For example, Geodia spp., Stelletta normani and Stryphnus ponderosus occur in full salinity, typically 34-35 psu (Klitgaard & Tendal, 2004; Kutti et al., 2013; Murillo et al., 2012). In modelling work undertaken by Knudby et al. (2013), Geodia sp. was hypothesized to have a minimum bottom salinity tolerance threshold within the range of 34.3-34.8 psu, but they acknowledged that further investigation was needed. Howell et al. (2016) similarly quote a narrow salinity range of 34.8-35.5 ppt for 'ostur' habitat.  In the Faroe-Shetland Channel, Kazanidis et al. (2019) observed that the transects with the highest densities of sponges had salinity values that ranged from 34.91 to 35.13 psu. Kazanidis et al. (2019) also found that salinity was a significant variable in explaining variations in sponge density. Beazley et al. (2013) similarly noted that salinity played a significant role in structure-forming sponge abundance. Evidence of direct salinity experimentation on deep-sea sponges is scarce but changes in salinity are not thought to induce spawning (Spetland et al., 2007).

Sensitivity assessment. Although changes in salinity are unlikely, the highly stable nature of water masses found at upper bathyal depths and the salinity range that the characterising species occur in suggests that the biotopes are likely to be intolerant of salinity changes. Therefore, if exposed to increased salinity, resistance is likely to be ‘Medium’, resilience is likely to be ‘Low’ and sensitivity would be considered as ‘Medium’. Please note that this assessment is made with low confidence due to the limited evidence on the effects.  

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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

Evidence

Changes in salinity are unlikely due to the depth at which the biotopes are found, combined with the distance from shore and the low potential for brine or freshwater discharge. However, salinity (along with other factors) is an important controller of modelled deep-sea sponge distribution (Howell et al., 2016; Knudby et al., 2013). Many studies examining the distribution of deep-sea sponges are limited to recording the conditions present at natural sponge aggregations. For example, Geodia spp., Stelletta normani and Stryphnus ponderosus occur in full salinity, typically 34-35 psu (Klitgaard & Tendal, 2004; Kutti et al., 2013; Murillo et al., 2012). In modelling work undertaken by Knudby et al. (2013), Geodia sp. was hypothesized to have a minimum bottom salinity tolerance threshold within the range of 34.3-34.8 psu, but they acknowledged that further investigation was needed. In the Faroe-Shetland Channel, Kazanidis et al. (2019) observed that the transects with the highest densities of sponges had salinity values that ranged from 34.91 to 35.13 psu. Kazanidis et al. (2019) also found that salinity was a significant variable in explaining variations in sponge density.  Evidence of direct salinity experimentation on deep-sea sponges is scarce but changes in salinity are not thought to induce spawning (Spetland et al., 2007).

Sensitivity assessment. Although changes in salinity are unlikely, the highly stable nature of water masses found at upper bathyal depths and the salinity range that the characterising species occur in suggests that the biotopes are likely to be intolerant of salinity changes. Therefore, if exposed to reduced salinity, resistance is likely to be ‘Medium’, resilience is likely to be ‘Low’ and sensitivity would be considered as ‘Medium’. Please note that this assessment is made with low confidence due to the limited evidence on the effects.     

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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

Evidence

As filter feeders, water flow (e.g., deep-sea bottom current) is important to the characterizing species for the supply of food and removal of metabolic waste. Studies into a range of deep-sea sponge species, including Pheronema carpenteri, Asconema sp. and the Geodiidae, have concluded that distribution and current flow may be linked, whether through supply of larval recruits, food, favourable dissolved oxygen conditions or removal of sediment (Beazley et al., 2013; Rice et al., 1990; White, 2003). For example, Roberts et al. (2018) stated that enhanced currents benefit sponge grounds in the Arctic (dominated by Geodia parva, Geodia hentscheli and Stelletta rhaphidiophora) by improving food and larval supply, as well as preventing smothering by the settling of suspended sediments.

The resistance of the characterizing species to changes in water flow regime is poorly understood. Results from modelling studies suggest that changes in water flow regime may not have a significant impact in some areas but have a larger impact on others. Modelling results from the north-west Atlantic suggested that deep-sea sponge grounds were found primarily in areas of high (>0.1 m/s) maximum bottom current flow and that minimum current speed was also an important variable in the model output, especially in slope waters (Knudby et al., 2013). In high-energy environments, Geodia barretti is found attached to stable substrata, while in areas with lower energy it is not attached but incorporates small stones in order to maintain negative buoyancy (Tjensvoll et al., 2013). This observation suggests that the species would not be negatively impacted by any changes in water flow.

Water flow conditions are vitally important for filter feeders, and to prevent excess sedimentation. While the characterizing sponge species are less likely to suffer damage in elevated currents due to their massive morphology, variable flow may reduce the flux of food. This, in turn, may reduce growth and increase metabolic demand through the additional pumping or restriction of larval dispersal. Provided there are no long-term physiological changes, the characterizing species should be able to revert to baseline levels of activity if changing water flow pressure was to cease.

Sensitivity assessment. A decrease at the benchmark level (0.1-0.2 m/s) is unlikely to have a significant effect on the normal high flow rates that this biotope experiences, and as the characterizing species normally occur in areas of higher flow rates (such as in the Faroe Shetland Channel; Masson et al., 2004), then an increase is unlikely to be detrimental. Therefore, resistance is assessed as ‘High’, resilience as ‘High’ and the biotope is considered ‘Not sensitive’ at the benchmark level.

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Emergence regime changes [Show more]

Emergence regime changes

Benchmark.  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

Evidence

The Geodia and other massive sponges biotopes are found at upper bathyal depths. Therefore, they will not be affected by changes in emergence regime and the biotopes are assessed as ‘Not relevant’.

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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

Evidence

The Geodia and other massive sponges biotopes are found at upper bathyal depths. Therefore, they will not be affected by changes in nearshore wave exposure and the biotopes are assessed as ‘Not relevant’.

Not relevant (NR)
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Not relevant (NR)
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Not relevant (NR)
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Chemical Pressures

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ResistanceResilienceSensitivity
Transition elements & organo-metal contamination [Show more]

Transition elements & organo-metal contamination

Benchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail

Evidence

This pressure is ‘Not assessed’, but any evidence is presented where available.

A study on the effects of trace metals (copper, iron, zinc, magnesium) on the shallow-water congeneric species Geodia cydonium in the Mediterranean, showed altered cell functions and effects on the immune system (Saby et al., 2009). Furthermore, all metals studied inhibited enzyme activity (Saby et al., 2009).  The congeneric shallow-water species Stelletta anancora was found to selectively accumulate the toxin arsenic, even if it was at normal environmental levels (Araújo et al., 2003, cited in Schoenberg, 2016).

Barite (BaSO4), a solid primary particulate compound in water-based drilling muds, can contain low concentrations of heavy metals and metalloid impurities (such as arsenic, chromium, copper, lead, nickel and zinc) (Neff, 2008, 2010). After Geodia barretti was exposed to barite for 14 days, the concentrations of the metals copper and strontium were not significantly different from that of sponges exposed to seawater (Edge et al., 2016). Total concentrations of lead, however, were significantly higher in sponges exposed to 30 mg/L of barite for 14 days, compared to those exposed to seawater. Nevertheless, the levels of lead were below those known to cause adverse effects in other sponge species (Cebrian et al., 2003).

Laboratory experiments were carried out to assess the impact of heavy metal contamination on the larval settlement and survival of the shallow-water sponge species Crambe crambe in the Mediterranean (Cebrian and Uriz, 2007). The results showed no negative effects of short term (1 week) exposure to copper and cadmium at concentrations of 30 µg/l and 5 µg/l, respectively. This study also concluded that the synergetic negative effect of copper and hydrocarbons on Crambe crambe decreases settlement further, which may cause a decline in sponge populations.

Fang et al. (2018) investigated the effects of suspended barite on Geodia barretti and Stryphnus fortis (similar to the characterizing species Stryphnus ponderosus). This study also looked at suspended natural sediment effects and bentonite (see ‘Change in suspended solids’ and ‘Introduction of other substances’ pressures). No sponge mortality was observed over the 33-day exposure periods, during which concentrations of ≤15.2 mg/l of barite were used. However, the sponges were covered by deposited particles. Spicule protrusion, one mechanism to reduce smothering by sediment, was observed for Geodia barretti, however, it was apparently not effective. Stryphnus fortis showed reduced tissue oxygenation upon exposure to barite, likely to be caused by reduced pumping, another mechanism to reduce smothering by sediment (Bell et al., 2015; Fang et al., 2018). Exposure to fine particles caused reduced oxygen consumption, plus correlated with reduced nitrite and nitrate release by the sponges. Geodia barretti recovered its oxygen-nitrogen metabolism (back to control levels) following abatement of bentonite after 33 days. After 33-days following abatement of barite, recovery did not occur for either of the barite impacted sponge species. Stryphnus fortis indicated possible long-term damage from barite, whereby the net ammonium flux displayed strong delayed responses during the recovery period (Fang et al., 2018).

Exposure to barite also induces cellular stress (lysosomal instability) in Geodia barretti (Edge et al., 2016). Lysosomal membrane stability (LMS) in Geodia barretti decreased with an increase in the concentration of suspended barite following 12-hour exposure, and levels of LMS were significantly lower with higher concentrations (50 mg and 100 mg per litre of total suspended solids (TSS/L); Edge et al., 2016). Longer-term and continuous exposure to barite for 14 days caused a decrease in LMS with increasing total suspended solids and length of exposure. Intermittent exposure to barite again resulted in lowered LMS with increases in TSS. Sponges exposed to barite at 30 mg TSS/L after six days had lower LMS than those exposed to reference sediment, but at 14 days the difference was not significant. LMS in sponges exposed to intermittent barite at 10 mg TSS/L was similar to reference sediments and seawater, indicating that no cellular stress is caused at this concentration of barite. Intermittent exposures, with 12-hour recovery periods, did not allow full recovery from cellular toxicity, indicating that longer recovery is required. However, the long-term effect of barite on LMS was reduced with this intermittent exposure. Overall, LMS levels in Geodia barretti were lowered by up to 30% with exposure to barite (Edge et al., 2016).

Not Assessed (NA)
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Not assessed (NA)
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Not assessed (NA)
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Hydrocarbon & PAH contamination [Show more]

Hydrocarbon & PAH contamination

Benchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail

Evidence

This pressure is ‘Not assessed’, but any evidence is presented where available.

Stévenne (2018) exposed Geodia barretti to a weathered blend of crude oil for eight days, at varying concentrations (33, 100 and 300 µg/l) to mimic conditions experienced after an oil-spill event. No sponge mortality, necrosis, tissue loss or colour change occurred during the experiment. Furthermore, no significant changes in respiration rates occurred, although a decrease of up to 52% was observed after 24 hours of exposure to oil compared to control sponges. Varying patterns of increased and decreased rates of respiration were observed over the exposure time and compared to the control. A higher mean abundance in destabilised lysosomal cells occurred with the increase in oil concentration. Respiration rates and lysosomal stability of all sponges from all treatments returned to levels comparable to the control after the recovery period of 30 days (Stévenne, 2018).

Laboratory experiments were carried out to assess the impact of polycyclic aromatic hydrocarbon (PAH) contamination on the larval settlement of the shallow-water sponge species Crambe crambe in the Mediterranean (Cebrian & Uriz, 2007). The results showed that exposure for 10 days inhibited the settlement of sponge larvae to a certain extent (25% and 30% in 500 and 1000 mg/l, compared to 65% in control). This study also showed that the synergetic negative effect of copper and hydrocarbons on Crambe crambe decreases settlement further, which may cause a decline in sponge populations.

Vad et al. (2017b) investigated the effect of hydrocarbons and dispersants on the shallow-water sponge species Halichondria panicea. During exposure to hydrocarbon contaminated seawater, the filtration behaviour of the sponge was diminished, and no recovery occurred during the 48-hour period after exposure.

Not Assessed (NA)
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Not assessed (NA)
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Not assessed (NA)
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Synthetic compound contamination [Show more]

Synthetic compound contamination

Benchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail

Evidence

This pressure is ‘Not assessed’, and no evidence was available regarding the effect of synthetic compound contamination on the Geodia and other massive sponges biotopes.

Not Assessed (NA)
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Not assessed (NA)
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Not assessed (NA)
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Radionuclide contamination [Show more]

Radionuclide contamination

Benchmark. An increase in 10µGy/h above background levels. Further detail

Evidence

No evidence

No evidence (NEv)
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Not relevant (NR)
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No evidence (NEv)
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Introduction of other substances [Show more]

Introduction of other substances

Benchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail

Evidence

This pressure is ‘Not assessed’, but any evidence is presented where available.

Fang et al. (2018) investigated the effects of suspended bentonite (mineralised volcanic ash) on Geodia barretti and Stryphnus fortis (similar to the characterizing species Stryphnus ponderosus). This study also looked suspended natural sediment and barite effects (see ‘Change in suspended solids’ and ‘Transition elements & organo-metal (e.g. TBT) contamination’ pressures). No sponge mortality was observed over the 33-day exposure periods, during which concentrations of ≤15.2 mg/l of bentonite were used, however, the sponges were covered by deposited particles. Spicule protrusion, one mechanism to reduce smothering by sediment, was observed for Geodia barretti, however, it was apparently not effective. The reduced tissue oxygenation in Geodia barretti and Stryphnus fortis upon exposure to bentonite was likely to be caused by reduced pumping, another mechanism to reduce smothering by sediment (Bell et al., 2015; Fang et al., 2018). Exposure to fine particles caused reduced oxygen consumption, plus correlated with reduced nitrite and nitrate release by the sponges. Geodia barretti recovered its oxygen-nitrogen metabolism (back to control levels) following abatement of bentonite after 33 days. Stryphnus fortis indicated possible long-term damage from bentonite, whereby the net ammonium flux displayed strong delayed responses during the recovery period (Fang et al., 2018). Exposure of Geodia barretti to suspended bentonite up to a total suspended solids (TSS) concentration of 100 mg TSS/L for 12 hours caused no significant change to lysosomal membrane stability (LMS) (Edge et al., 2016).

Not Assessed (NA)
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Not assessed (NA)
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Not assessed (NA)
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De-oxygenation [Show more]

De-oxygenation

Benchmark. 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

Evidence

Deoxygenation is known to cause mortality in a range of benthic organisms, especially sessile taxa that cannot escape unfavourable conditions. For the characterizing species Geodia barretti, there is some evidence of behavioural adaptations (reduced pumping and respiration rates) that allow the sponges to restrict oxygen consumption as a protective mechanism when exposed to increased sediment load, which can cause short-term depletion of oxygen (Kutti et al., 2015; Schoenberg, 2016; Tjensvoll et al., 2013). There is also evidence of natural hypoxia and suboxia in some Demospongiae taxa, related to internal microbial conditions (Lavy et al., 2016). However, Leys et al. (2018) suggested that changes in oxygenation may negatively affect Geodia barretti.  Leys et al. (2004) found that Hexactinellida (glass sponges) were rare (fewer than eight live sponges per 10 m2) in regions of fjords in British Columbia where dissolved oxygen levels fell below 2 ml/l. It may be possible that Demospongiae share a similar pattern, however, no evidence for this class could be found. As there is no direct evidence on the resistance or resilience of the characterizing species in a reduced oxygen scenario at benchmark pressure, this pressure is assessed as ‘No evidence’.

No evidence (NEv)
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Not relevant (NR)
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No evidence (NEv)
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Nutrient enrichment [Show more]

Nutrient enrichment

Benchmark. Compliance with WFD criteria for good status. Further detail

Evidence

There is some evidence to suggest that nutrient enrichment may be beneficial to sponges by providing additional food resources. Leys et al. (2018) provided the first direct in vivo measurement of nutrient flux in Geodia barretti. Ammonium and nitrite are taken up by Geodia barretti, and nitrate is produced (Leys et al., 2018). Silicate is also required by demosponges (including Geodia sp., Stryphnus ponderosus and Stelletta normani) for biomineralization of spicules to produce their skeletons, which shapes the sponge growth, and allows cell organization and development of the aquiferous system (Scanes et al., 2018; Uriz et al., 2003). However, there is no evidence available to assess the effect of nutrient enrichment on the characterizing species at benchmark pressure. Nevertheless, by definition, the biotopes are considered to be 'Not sensitive' at the pressure benchmark, which assumes compliance with good status as defined by the WFD.

Not relevant (NR)
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Not relevant (NR)
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Not sensitive
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Organic enrichment [Show more]

Organic enrichment

Benchmark. A deposit of 100 gC/m2/yr. Further detail

Evidence

It is possible that a deposit of organic matter will provide additional food resources for the filter-feeding characterizing species, although Leys et al. (2018) found that little of the carbon in Geodia barretti comes from detritus, despite requiring it for energy. Howell et al. (2016) further suggest that the amount of particulate organic carbon, on which sponges feed, is a key driver of 'ostur' sponge distribution. In two Scandinavian fjords, the reproduction of Geodia barretti is known to be linked to the sedimentation of particulate organic matter (Spetland et al., 2007).  Gamete release occurs in early summer and October when sedimentation of particulate organic matter is at its highest, after the phytoplankton blooms in spring and autumn (Spetland et al., 2007). This suggests that the reproduction of Geodia barretti is highly dependent on the seasonality of food supplies (Hogg et al., 2010).

Sensitivity assessment. Geodia barretti requires organic matter in suspension to filter feed (Howell et al., 2016) and the seasonality of particulate organic matter is known to play a role in the reproduction pattern of the species (Hogg et al., 2010; Spetland et al., 2007). Therefore, the species would likely benefit from any organic enrichment, so long as it does not negatively disrupt the normal reproduction cycle of the species. Therefore, resistance for the organic enrichment pressure is assessed as ‘High’, resilience as ‘High’, and the biotopes are considered ‘Not sensitive’ at the benchmark level.

High
Medium
Medium
Medium
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High
High
High
High
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Not sensitive
Medium
Medium
Medium
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Physical Pressures

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ResistanceResilienceSensitivity
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

Evidence

All marine habitats and benthic species are considered to have a resistance of ‘None’ to this pressure and to be unable to recover from a permanent loss of available habitat (resilience is ‘Very low’). The Geodia and other massive sponges biotopes are therefore considered to have ‘High’ sensitivity to this pressure.

None
High
High
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Very Low
High
High
High
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High
High
High
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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

Evidence

In the Faroe-Shetland Channel, Kazanidis et al. (2019) found that the highest densities of Category 3 sponges (i.e., massive, spherical or papillate, including Geodia spp.) were associated with substrata consisting of cobbles with boulders. Kazanidis et al. (2019) also found that the substratum (specifically cobbles with boulders) was a significant variable in explaining variations in sponge density. Furthermore, there is some evidence suggesting that Geodia spp. larvae can settle/grow on hard substrata (cobbles and boulders) (Freese, 2001; Henry and Roberts, 2014). However, previous surveys found the characterizing species associated with sedimentary habitats only (Klitgaard & Tendal, 2004).

Sensitivity assessment. Nonetheless, a change from sedimentary to hard rock substratum is likely to impact the characterizing species and the physical habitat. As a specific sediment type defines biotopes, a change in sediment type will result in a change in the biotope classification and therefore the loss of the original biotope. Hence, resistance is assessed as 'None'. As this pressure is considered as a permanent change, resilience is assessed as 'Very Low', and sensitivity is, therefore, assessed as 'High'.

None
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Very Low
High
High
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High
High
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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

Evidence

Although the Geodia and other massive sponges biotopes are found in both coarse and mixed sediment habitats (Parry et al., 2015) it is possible that changes in the Folk classification may have a small impact on the characterizing species. While a change to mixed sediment or coarse sediment may not have significant implications for the characterizing species, there is some evidence that changes to finer substrata i.e. sand or muddy sand will be detrimental. The distribution of this sponge community is strongly connected with the presence of pebbles and cobbles within the substratum (e.g. Kazanidis et al., 2019). Therefore, the species would not have a cobble to “hold on” to for anchorage on sandy substrata. Furthermore, on muddy sediment with strong bottom currents, the resuspension of the sediment fine fraction could clog the feeding capability of the filter feeder sponges. Klitgaard & Tendal (2004) found that silty or muddy sediments of volcanic origin were not suitable substrata for deep-sea sponges. Boreal ‘ostur’ assemblages have also been shown to increase the physical heterogeneity of the bottom, changing the local sediment structure over time through the release of spicules after death, resulting in muddy spicule mats covering the bottom (Klitgaard & Tendal, 2004).

Sensitivity assessment. A change in sediment type will result in a change in the biotope classification and, therefore, the loss of the original biotope. Although Geodia can survive in mixed or coarse sediments, sand or muddy-sand is not suitable, so resistance is assessed as ‘None’. As this pressure is considered as a permanent change, resilience is assessed as 'Very Low', and the biotope is considered to have ‘High’ sensitivity to a change in seabed type by one Folk class.

None
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Very Low
High
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High
High
High
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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

Evidence

As the Geodia and other massive sponges biotopes are characterized by sessile invertebrates, removal of the substratum at the benchmark level would cause the destruction of the biotopes within the affected area. Recovery would be severely impacted given the slow growth rate of the characteristic species, and as Geodia barretti is an annual spawner with only a short-range larval dispersal and high larval retention (Mariani et al., 2006), resulting in low connectivity (Knudby et al., 2013).

Sensitivity assessment.  As the upper 30 cm of sediment is the most important fraction for accompanying fauna, nutrient recycling, etc., resistance is assessed as 'None'. Resilience is likely to be 'Very low’ based upon the slow growth rates of Geodia sp. (i.e., several decades; Klitgaard and Tendal, 2004), likely extreme lifespans (Fallon et al., 2010; Leys and Lauzon, 1998), and low connectivity (i.e., short-range larval dispersal and high larval retention; Knudby et al., 2013). Therefore, sensitivity is assessed as ‘High’.

None
Low
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Very Low
High
Medium
High
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High
Low
Low
Low
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Abrasion / disturbance of the surface of the substratum or seabed [Show more]

Abrasion / disturbance of the surface of the substratum or seabed

Benchmark. Damage to surface features (e.g. species and physical structures within the habitat). Further detail

Evidence

Species of Demospongiae may be more resistant to surface abrasion than Hexactinellid species (glass sponges) due to their proteinaceous skeletons that make their structure more flexible (Serpetti et al., 2014). However, Freese (2001) found that Geodia sp. broke or were turned over when nudged by a submersible in their study. Observations from northern Norway show that trawl gear can transport whole sponge individuals (such as Geodia spp. and Stelletta sp.) and deposit them in lines when the gear is recovered, or in dips in seabed topography (Buhl-Mortensen et al., 2016). Kędra et al. (2017) mentioned that some large sponges were observed in old trawling tracks in the Barents Sea, and suggested that these were likely dislocated and moved by fishing activities.

In the Barents Sea, Kędra et al. (2017) found that, in a location where both Geodia barretti and Geodia macandrewii occur, the mean individual sponge biomass was higher in untrawled areas. In trawled areas, sponges were more than five times less frequent (and 18 times less frequent in one study area) compared to untrawled areas. Geodia barretti and Geodia macandrewii were found at both the trawled and untrawled sites in one study area with low fishing pressure. However, in the other two study areas, at higher fishing intensity, a strong reduction in the biomass of Geodia barretti was recorded compared to untrawled sites in the same areas. For example, in one area, sponge mean individual biomass was 32.2 g m-2 (± 57.0) in untrawled sites, and 19.1 g m-2 (± 54.7) in trawled sites. It is important, however, to note the very high variability in biomass recorded. In the Faroe-Shetland Channel, Kazanidis et al. (2019) found that demersal fisheries pressure was the most significant variable in explaining variations in sponge density. The higher densities of sponges were found in the area with the lowest values of demersal landings. Densities in the areas with higher demersal landings were significantly lower.

Geodia spp. are noted to be able to survive damage from physical contact with trawl gear, as well as to survive being handled on ships, and they can re-establish on the seabed once returned (Jørgensen et al., 2016). Sponges left damaged by trawls are vulnerable, however, particularly as their filter-feeding systems may be clogged by non-food particles (Løkkeborg and Fosså, 2011, cited in Jørgensen et al., 2016). Geodia barretti explants are able to continue their reproductive cycle following manipulation (Hoffmann et al., 2003), and tissue regeneration is known to occur in sponges (albeit not specifically Geodia sp.), so Jørgensen et al. (2016) suggest that trawl-induced mortality is equivocal.

Teixidó et al. (2004) conducted an assessment in the Weddell Sea, looking at the re-colonization of the benthic ecosystem, including Demospongiae, after iceberg disturbance. Re-colonization was assessed over three stages based on qualitative criteria (e.g., faunal composition, categorical abundance and seabed relief features). Although no quantitative studies have been carried out to characterize the time frame of these patterns, Demospongiae species appeared to be quicker at recolonizing impacted mobile substrata than Hexactinellid species (glass sponges), but massive sponges defined the late stages of recovery.

Freese (2001) assessed the recovery of Geodia sp. (later re-identified as likely Poecillastra tenuilaminaris of the same sub-order, Astrophorina; Malecha & Heifetz, 2017) in the Gulf of Alaska (at 206-209 m depth) following experimental trawling (one trawl per area). Furrows in the substratum from the trawling activity, and gouges from the dragging of boulders, were found to still be prominent after one year, with little evidence of backfilling. No new colonization was evident one year after the trawling and, of the surviving Geodia sp. individuals, 56.3% were found lying on the bottom either torn or intact, and 40.6% were undamaged. The remaining 3.1% were upright but torn. Geodia spp. did not show any sign of necrosis and, although damaged sponges did not show signs of repair or regrowth, most of the knocked over sponges or pieces of sponge that had been torn off still appeared viable (Freese, 2001).

In a follow-up study, 13 years post-trawling, Malecha & Heifetz (2017) found that the average density of Poecillastra tenuilaminaris (re-identified from Geodia sp.) was nearly equal between trawl and reference transects.  However, video analysis showed that 19.3% of the sponges in trawl transects were damaged (either lying on the bottom or torn). This was twice the rate of that in reference transects, and this species had the highest percentage of missing tissue compared to other species. On detailed observation, 57.1% were damaged in trawled areas versus 16.7% in reference transects. Seafloor gouging, boulder displacement and damaged sponges were still visible on some trawl transects after 13-years.    

A modelling study also found that if a sponge community in the Aleutian Islands, Alaska, which included Geodiidae, was subjected to bottom trawling, it would likely only recover to 80% of its original biomass within a 20-year timeframe, in the absence of further damage/removal (Rooper et al., 2011). Care should be taken however when using the results of models outside of a proper validation (Rooper et al., 2018). 

As mixed sediment biotopes can contain fractions of muddy-sand, surface abrasion can also cause resuspension of these sediments, in addition to mechanical damage, which may remove the top layer of sediment. Abrasion pressure can, therefore, cause the release of anoxic sediments or nutrients, and potentially siltation of benthic fauna (Schoenberg, 2016).  These pressures are assessed separately (see ‘Changes in suspended solids’, ‘Smothering and siltation changes’ and ‘Nutrient enrichment’). 

Sensitivity assessment. Although some Geodia spp. are reported to survive damage from physical contact with trawl gear, the benchmark level of pressure states ‘damage to seabed surface features’. At this benchmark level, there is significant evidence to support effects occurring, and the top layer of the sediment is likely to be removed. As such, the community and associated habitat of the biotopes will be impacted, so resistance is assessed as ‘None’, resilience as ‘Very low’ and overall sensitivity as ‘High’. 

None
High
High
High
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Very Low
High
Medium
High
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High
High
Medium
High
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Penetration or disturbance of the substratum subsurface [Show more]

Penetration or disturbance of the substratum subsurface

Benchmark. Damage to sub-surface features (e.g. species and physical structures within the habitat). Further detail

Evidence

Penetration and or disturbance of the substratum would result in similar, if not identical effects as the Abrasion pressure.

Sensitivity assessment. Although some Geodia spp. are noted to be able to survive damage from physical impacts, since the biotopes are characterized by sessile invertebrates that live on the surface of the sediment, damage to the sub-surface seabed would likely mean that most of the population will be killed, damaged or removed by the pressure. This would result in the loss of the biotopes, and re-colonization will have to come from other areas (Serpetti et al., 2014). Furthermore, penetration is likely to cause the removal of the top layer of sediment. As such, the community and associated habitat of the biotopes will be impacted, so resistance is assessed as ‘None’, resilience as ‘Very low’ and sensitivity as ‘High’.

None
High
High
High
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Very Low
High
Medium
High
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High
High
Medium
High
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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

Evidence

Laboratory studies have shown severe reductions in respiration rates of Geodia barretti when subjected to high levels of suspended sediments for four hours (Tjensvoll et al., 2013). The study found decreases of 52% (at 50 mg/l), 86% (at 100 mg/l) and 67% (at 500 mg/l). However,  specimens returned to a baseline metabolic rate four hours after exposure (Tjensvoll et al., 2013). In the higher sediment loads, sponges were noted to be actively pumping one hour after the exposure period ended. In another study of four-hour exposure to suspended crushed rock particles at 500 mg/l, Kutti et al. (2015) similarly found that the oxygen consumption of Geodia barretti decreased significantly (50%) from pre-exposure consumption. However, all sponges recovered within the 30-minute flushing cycle, and oxygen consumption did not differ significantly from the pre-exposure rates.

In long-term exposure (50 days) of Geodia barretti to high (50 mg/l) and low (10 mg/l) concentrations, oxygen consumption was 50% lower in sponges exposed to crushed rock compared to those exposed to natural bottom sediments (Kutti et al., 2015). However, after 29 days of cyclic exposure, the sponges’ respiration rates did not return to that of unexposed sponges and were reduced by >60% during the daily 12-hour recovery interval (Kutti et al., 2015). This suggested that a recovery time of more than 12 hours is required. The reductions in oxygen consumption were a result of a decrease or arrest in pumping, which was a response to high suspended solid concentrations (Tjensvoll et al., 2013). Bell et al. (2015) also suggested that the suppression of oxygen consumption may be a defence mechanism to prevent the internal sponge tissue being exposed to sediment. These studies, therefore, indicate that Geodia barretti has well developed responses to cope with short periods of elevated concentrations of suspended natural bottom sediments. Tjensvoll et al. (2013) also found that exposure of Geodia barretti to low sediment concentrations of 10 mg/l had little or no effect, although there were some variations in respiration rate during the recovery period.

Fang et al. (2018) investigated the effects of suspended natural sediment (and barite and bentonite) on Geodia barretti and Stryphnus fortis (similar to the characterizing species Stryphnus ponderosus). No sponge mortality was observed over the 33-day exposure periods, at concentrations of ≤15.2 mg/l, however, the sponges were covered by deposited particles. Reduced tissue oxygenation occurred in Stryphnus fortis on exposure to suspended natural sediments and this was likely to be caused by reduced pumping (Bell et al., 2015; Fang et al., 2018). Exposure to fine particles caused reduced oxygen consumption. It is suggested that finer particles (<63 µm) may have greater effects than larger particles (Kutti et al., 2015) on the metabolism in Geodia barretti, even at 10 mg/l (Fang et al., 2018), but Stryphnus fortis samples were less sensitive. However, Geodia barretti recovered its oxygen-nitrogen metabolism (to control levels) following abatement of suspended natural sediment after 33 days. However, net ammonium flux displayed strong delayed responses during the recovery period in Stryphnus fortis, which indicates possible long-term damage from suspended natural sediment.

In a further study, exposure of Geodia barretti to suspended natural (reference) sediment up to a total suspended solids concentration of 100 mg/l for 12 hours caused no significant change to lysosomal membrane stability (LMS) (an indication of cellular stress) (Edge et al., 2016). Furthermore, there was no significant difference in LMS in sponges exposed to seawater compared to suspended reference sediments at 10, 50 or 100 mg/l for 12 hours. Over a slightly longer exposure, sponges exposed to reference sediment at 30 mg/l after six days did show lower LMS and lower total organic energy than those exposed to seawater, but at 14 days there were no significant differences. The increase in lysosomal stability with length of exposure suggested possible acclimatisation in Geodia barretti at <30 mg/l (Edge et al., 2016 cited in Scanes et al., 2018). A decrease in respiration levels was also recorded when Geodia atlantica was exposed to suspended sediments at 10 mg/l over a period of 40 days (Scanes et al., 2018). Furthermore, lower silicate uptake (used for biomineralization to produce their skeletons) occurred after 33 and 40 days of exposure to suspended sediments (Scanes et al., 2018). After 26 days of exposure, lysosomal membrane instability was increased in comparison to the control, but this effect was not observed again at 33 or 40 days, which again suggests that Geodia atlantica can acclimatise after 26 days of exposure to low levels of suspended sediment (Scanes et al., 2018).

Sensitivity assessment. At the benchmark level of change in one rank on the WFD scale for one year (i.e. from 10 to 100 mg/l), suspended solids may reduce the survivability of the characterizing species in the long term, despite there being some evidence of acclimatisation (Scanes et al., 2018). The effects also vary with the type and particle size of the suspended sediment. Higher levels of sediment exposure are likely to also increase the recovery period (Pineda et al., 2017b). Therefore, resistance is assessed as ‘Medium’, resilience as ‘Low’ and overall sensitivity is assessed as ‘Medium’.

Medium
High
High
High
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Low
High
Medium
High
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Medium
High
Medium
High
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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

Evidence

Deep-sea sponge aggregations occur in high water flow environments, which will affect how long any sedimentation deposits remain, and whether this is long enough to have any significant effect. Roberts et al. (2018) mention that enhanced currents at sponge grounds prevent smothering by settling suspended sediments. Jones et al. (2012) noted that drill cuttings were removed over time in the Faroe-Shetland Channel by lateral advection during high current events of velocities up to 1 m/s (Masson et al., 2004) and from bioturbation resulting in vertical redistribution in the sediments. Deposition of up to 1.5 m of drill cuttings in the Faroe-Shetland Channel resulted in a reduction of Geodia spp. from 13.3 individuals per 100 m2 before the impact, to 0.3 individuals per 100 m2 10 years later (Jones et al., 2012). After three years, Jones et al. (2012) noted that a significant amount of drill cuttings had been removed, particularly in areas of lower initial deposition. The pressure used in this study, however (up to 1.5 m of sediment cover) far exceeds the benchmark level (up to 5 cm of fine material). Drill cuttings changed the sediment median grain size from coarse to fine. They are also known to contain compounds that cause ecotoxicological effects, e.g. from added mud chemicals or from impurities in the weight material (Trannum et al., 2010).  Therefore, the contaminants may have increased the impact and/or delayed the recovery of Geodia spp. For more information on the effects of specific drilling mud substances (bentonite and barite), please see the ‘Introduction of other substances’ pressure.

Fang et al. (2018) observed that Geodia barretti and Stryphnus fortis (similar to the characterizing species Stryphnus ponderosus) were covered by deposited particles following a 33-day exposure to suspended solids (≤15.2 mg/l). Spicule protrusion, one mechanism to reduce smothering by sediment, was observed for Geodia barretti, however, it was apparently not effective to the intensive sediment deposition the sponges were subject to in the study.  Another mechanism to reduce smothering by sediment in sponges is to reduce pumping and, therefore, reduce tissue oxygenation. Reduced pumping was observed in Stryphnus fortis on exposure to suspended natural sediments (Bell et al., 2015; Fang et al., 2018). In another experiment on Geodia barretti, images of the sponges before and after the exposure to suspended particles showed ‘light’ deposition of sediment on the sponge surface (Kutti et al., 2015).

The effects of burial on sponges, in general, were discussed by Schoenberg (2016), who suggested that resistance may be influenced by the nature of the deposited sediment and duration of burial. It was hypothesised that coarse material may reduce the amount of deoxygenation that the sponge experiences from burial. It is also expected that sponges may be able to survive temporary burial (Hoffmann et al., 2008) but would commonly die if left buried (Schoenberg, 2016). Resistance may vary by species and be dependent on the morphology and where individuals survive, there is likely to be a large metabolic cost in clearing the sponge’s respiratory channels (Schoenberg, 2016). In addition, Schoenberg (2016) reported a range of concerns surrounding siltation on sponges in general, including behavioural adaptations, mucus production, and phagocytosis of ingested material.

Sensitivity assessment. While discrete siltation events may not be detrimental to the characterizing species, the nature of the deposited material and the duration of burial may influence the sensitivity of the biotope. Geodia barretti and Stryphnus fortis (similar to the characterizing species Stryphnus ponderosus) both exhibit mechanisms to reduce smothering by sediments. However, high rates of sediment deposition (e.g. Fang et al., 2018; Jones et al., 2012) will cause the porous sponges to become clogged with sediment, resulting in reduced feeding capability (Klitgaard & Tendal, 2004). Mortality and low recovery rates (>10 years; Jones et al., 2017) can then occur, even after a significant amount of the sediment is removed (e.g. after three years, due to the high energy environments associated with the biotopes; Jones et al., 2017). Therefore, resistance is assessed as ‘Medium’, resilience as ‘Low’, and sensitivity as ‘Medium’. Any abrasion effects associated with the removal of this pressure are covered under the specific abrasion pressure.

Medium
High
High
High
Help
Low
High
Medium
High
Help
Medium
High
Medium
High
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

Evidence

Deep-sea sponge aggregations occur in high water flow environments, which will affect how long any sedimentation deposits remain, and whether this is long enough to have any significant effect. Roberts et al. (2018) mention that enhanced currents at sponge grounds prevent smothering by settling suspended sediments. Jones et al. (2012) noted that drill cuttings were removed over time in the Faroe-Shetland Channel by lateral advection during high current events of velocities up to 1 m/s (Masson et al., 2004) and from bioturbation resulting in vertical redistribution in the sediments. Deposition of up to 1.5 m of drill cuttings in the Faroe-Shetland Channel resulted in a reduction of Geodia spp. from 13.3 individuals per 100 m2 before the impact, to 0.3 individuals per 100 m2 10 years later (Jones et al., 2012). After three years, Jones et al. (2012) noted that a significant amount of drill cuttings had been removed, particularly in areas of lower initial deposition. The pressure used in this study, however (up to 1.5 m of sediment cover) far exceeds the benchmark level (up to 5 cm of fine material). Drill cuttings changed the sediment median grain size from coarse to fine. They are also known to contain compounds that cause ecotoxicological effects, e.g. from added mud chemicals or from impurities in the weight material (Trannum et al., 2010).  Therefore, the contaminants may have increased the impact and/or delayed the recovery of Geodia spp. For more information on the effects of specific drilling mud substances (bentonite and barite), please see the ‘Introduction of other substances’ pressure.

Fang et al. (2018) observed that Geodia barretti and Stryphnus fortis (similar to the characterizing species Stryphnus ponderosus) were covered by deposited particles following a 33-day exposure to suspended solids (≤15.2 mg/l). Spicule protrusion, one mechanism to reduce smothering by sediment, was observed for Geodia barretti, however, it was apparently not effective to the intensive sediment deposition the sponges were subject to in the study.  Another mechanism to reduce smothering by sediment in sponges is to reduce pumping and, therefore, reduce tissue oxygenation. Reduced pumping was observed in Stryphnus fortis on exposure to suspended natural sediments (Bell et al., 2015; Fang et al., 2018). In another experiment on Geodia barretti, images of the sponges before and after the exposure to suspended particles showed ‘light’ deposition of sediment on the sponge surface (Kutti et al., 2015).

The effects of burial on sponges, in general, were discussed by Schoenberg (2016), who suggested that resistance may be influenced by the nature of the deposited sediment and duration of burial. It was hypothesised that coarse material may reduce the amount of deoxygenation that the sponge experiences from burial. It is also expected that sponges may be able to survive temporary burial (Hoffmann et al., 2008) but would commonly die if left buried (Schoenberg, 2016). Resistance may vary by species and be dependent on the morphology and where individuals survive, there is likely to be a large metabolic cost in clearing the sponge’s respiratory channels (Schoenberg, 2016). In addition, Schoenberg (2016) reported a range of concerns surrounding siltation on sponges in general, including behavioural adaptations, mucus production, and phagocytosis of ingested material.

It is worth considering that currents and water flow will vary by location and as such, it is difficult to generalise about the effect of burial on the biotope. Nevertheless, benchmark levels of deposition are likely to represent a significant pressure to the biotope.

Sensitivity assessment. While discrete siltation events may not be detrimental to the characterizing species, the nature of the deposited material and the duration of burial may influence the sensitivity of the biotopes. Geodia barretti and Stryphnus fortis (similar to the characterizing species Stryphnus ponderosus) both exhibit mechanisms to reduce smothering by sediments. However, high rates of sediment deposition (e.g. Fang et al., 2018; Jones et al., 2012) will cause the porous sponges to become clogged with sediment, resulting in reduced feeding capability (Klitgaard & Tendal, 2004). Mortality and low recovery rates (>10 years; Jones et al., 2017) can then occur, even after a significant amount of the sediment is removed (e.g., after three years, due to the high energy environments associated with the biotopes; Jones et al., 2017). In areas of higher deposition, the removal of sediment, and therefore recovery, will take longer. As such, resistance is assessed as ‘Low’, resilience as ‘Very low’, and sensitivity is assessed as ‘High’. Any abrasion effects associated with the removal of this pressure are covered under the specific Abrasion pressure.

Low
High
High
High
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Very Low
High
Medium
High
Help
High
High
Medium
High
Help
Litter [Show more]

Litter

Benchmark. The introduction of man-made objects able to cause physical harm (surface, water column, seafloor or strandline). Further detail

Evidence

Although a number of studies (e.g., Chapron et al., 2018; Courtene-Jones et al., 2019, 2017; La Beur et al., 2019) have shown that microplastics are ingested by deep-sea invertebrates, the effects of the pressure are poorly understood, and no evidence could be found on the effects of litter on the characterizing species or other deep-sea sponges. This pressure is, therefore ‘Not assessed’.

Not Assessed (NA)
NR
NR
NR
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Not assessed (NA)
NR
NR
NR
Help
Not assessed (NA)
NR
NR
NR
Help
Electromagnetic changes [Show more]

Electromagnetic changes

Benchmark. A local electric field of 1 V/m or a local magnetic field of 10 µT. Further detail

Evidence

No evidence could be found on the effects of Electromagnetic changes on the characterizing species. Therefore, this pressure is assessed as ‘No evidence’.

No evidence (NEv)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
No evidence (NEv)
NR
NR
NR
Help
Underwater noise changes [Show more]

Underwater noise changes

Benchmark. MSFD indicator levels (SEL or peak SPL) exceeded for 20% of days in a calendar year. Further detail

Evidence

The Geodia and other massive sponges biotopes are characterized by invertebrates with no means to detect noise or vibration and, therefore, it is unlikely to be affected by changes in underwater noise. This pressure is assessed as ‘Not relevant’.

Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Introduction of light or shading [Show more]

Introduction of light or shading

Benchmark. A change in incident light via anthropogenic means. Further detail

Evidence

The Geodia and other massive sponges biotopes are characterized by invertebrates with no means to detect light, and they occur at considerable depth where little or no incident light penetrates from the surface. As such, the biotopes will not be affected by changes in light regime and this pressure is assessed as ‘Not relevant’.

Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Barrier to species movement [Show more]

Barrier to species movement

Benchmark. A permanent or temporary barrier to species movement over ≥50% of water body width or a 10% change in tidal excursion. Further detail

Evidence

The Geodia and other massive sponges biotopes are characterized by sessile invertebrates, which are not dependent on larvae from outside the population due to their short-range larval dispersal and high larval retention (Mariani et al., 2006), resulting in low connectivity (Knudby et al., 2013). Hence, these biotopes are unlikely to be affected by 'Barriers to species movement'. This pressure is assessed as ‘Not relevant’.

Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Death or injury by collision [Show more]

Death or injury by collision

Benchmark. 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

Evidence

The Geodia and other massive sponges biotopes are characterized by sessile invertebrates and are unlikely to be affected by an increased risk of collision defined under the pressure. This pressure is assessed as ‘Not relevant’.

Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Visual disturbance [Show more]

Visual disturbance

Benchmark. The daily duration of transient visual cues exceeds 10% of the period of site occupancy by the feature. Further detail

Evidence

The Geodia and other massive sponges biotopes are characterized by invertebrates that are not reliant on vision, and as such will not be affected by Visual disturbance. This pressure is assessed as ‘Not relevant’.

Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help

Biological Pressures

Use [show more] / [show less] to open/close text displayed

ResistanceResilienceSensitivity
Genetic modification & translocation of indigenous species [Show more]

Genetic modification & translocation of indigenous species

Benchmark. 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

Evidence

None of the characterizing species associated with the Geodia and other massive sponges biotopes are subject to cultivation or translocation, so this pressure is considered ‘Not relevant’.

Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Introduction or spread of invasive non-indigenous species [Show more]

Introduction or spread of invasive non-indigenous species

Benchmark. The introduction of one or more invasive non-indigenous species (INIS). Further detail

Evidence

No evidence could be found regarding the effect the 'Introduction or spread of non-indigenous species' on the Geodia and other massive sponges biotopes. Hence, the pressure is assessed as ‘No evidence’.

No evidence (NEv)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
Help
No evidence (NEv)
NR
NR
NR
Help
Introduction of microbial pathogens [Show more]

Introduction of microbial pathogens

Benchmark. 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

Evidence

While there is published evidence on immunity and the spread of diseases in sponges (Pita et al., 2018), evidence of disease in deep-sea sponges is scant. Evidence of natural defences against Vibrio harveyi (Isnansetyo et al., 2009) and natural antifoulants (Sjögren et al., 2011) has been shown in Geodia spp.. Luter et al. (2017) assessed the prevalence of disease affecting Geodia barretti in Norway. Within a total area of 534 m2 at depths from 100-220 m, 0-20% of the Geodia barretti community showed signs of disease, evident by brown/black discolouration on the surface of the sponge and extensive tissue integration and fouling. On investigation, distinctly different microbial communities were present in diseased sponges compared to healthy and non-diseased sponges. However, as the effects of this disease on the sponge function is not clear, this pressure is assessed as ‘No evidence’.

No evidence (NEv)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
No evidence (NEv)
NR
NR
NR
Help
Removal of target species [Show more]

Removal of target species

Benchmark. Removal of species targeted by fishery, shellfishery or harvesting at a commercial or recreational scale. Further detail

Evidence

The characterizing species associated with the Geodia and other massive sponges biotopes are not commercially targeted, hence this pressure is assessed as ‘Not relevant’.

Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Removal of non-target species [Show more]

Removal of non-target species

Benchmark. Removal of features or incidental non-targeted catch (by-catch) through targeted fishery, shellfishery or harvesting at a commercial or recreational scale. Further detail

Evidence

This pressure only assesses the biological effects of the removal of species. For evidence on the effects of the removal itself, please see the 'Physical damage' pressures.

Specimens of Geodia barretti, Geodia macandrewii, Geodia atlantica and Stryphnus ponderosus have been reported to exceed 70 cm in diameter and 24 kg wet weight (Klitgaard & Tendal, 2004), with other specimens of Geodia barretti having a wet weight of 36.7 kg (Leys et al., 2018). These large sizes and the massive sponge morphology of these characterizing species means that they sit raised off the seafloor, putting them at higher risk of being removed as incidental by-catch (Jørgensen et al., 2016). In northern Norway, whole sponge individuals (such as Geodia spp. and Stelletta sp.) were found deposited in lines (i.e. from when trawl gear is recovered) or in dips in the seabed topography (Buhl-Mortensen et al., 2016). Kędra et al. (2017) also observed large sponges in old trawling tracks in the Barents Sea, suggesting that these were likely dislocated and moved by fishing activities. If a significant proportion of the Geodia species are removed, the biological structure will be altered, and the habitat complexity reduced.

The degree of recovery would be reliant upon surviving sponges providing a nucleus for population recovery, as Geodia barretti only has a short-range larval dispersal and high larval retention (Mariani et al., 2006), resulting in low connectivity (Knudby et al., 2013). Freese (2001) noted that most knocked over Geodia sp., or pieces of Geodia sp. that had been torn off (following an experimental trawl) still appeared viable after one year, showing that even dislodged individuals can survive and support the recovery of the biotopes. After 13 years, the average density of Poecillastra tenuilaminaris (re-identified from Geodia sp.) was nearly equal between trawl and reference transects, despite sponges in the trawled areas still showing damage (twice that of sponges in reference transects) and having the highest percentage of tissue missing (Malecha & Heifetz, 2017). After eight months of cultivation in the field and adaptation in a laboratory, explants of Geodia barretti, which started at 4 x 4 x 4 cm in size, have pumping and respiration rates similar to that of full-grown individuals (Kutti et al., 2015). This time-period allowed cortex regeneration to take place (Hoffmann et al., 2003), although the survival rate during cultivation was only 50% (Kutti et al., 2015).

Sensitivity assessment. Removal of a large percentage of the characterizing species would alter the character of the biotope. The resistance to removal is assessed as ‘Low’ due to the inability of these species to evade collection. The resilience is assessed as ‘Low’, with recovery only being able to begin when the pressure is removed altogether. Therefore, the sensitivity is assessed as ‘High’.

Low
High
High
High
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Low
High
Medium
High
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High
High
Medium
High
Help

Bibliography

  1. Beazley, L.I., Kenchington, E.L., Murillo, F.J. & Sacau, M.D.M., 2013. Deep-sea sponge grounds enhance diversity and abundance of epibenthic megafauna in the Northwest Atlantic. ICES Journal of Marine Science, 70 (7), 1471-1490. DOI https://doi.org/10.1093/icesjms/fst124
  2. Bell, J.J., 2008. The functional roles of marine sponges. Estuarine, Coastal and Shelf Science, 79 (3), 341-353. DOI https://doi.org/10.1016/j.ecss.2008.05.002
  3. Bell, J.J., McGrath, E., Biggerstaff, A., Bates, T., Bennett, H., Marlow, J. & Shaffer, M., 2015. Sediment impacts on marine sponges. Marine Pollution Bulletin, 94 (1), 5-13. https://doi.org/10.1016/j.marpolbul.2015.03.030

  4. Buhl-Mortensen, L., Ellingsen, K.E., Buhl-Mortensen, P., Skaar, K.L. & Gonzalez-Mirelis, G., 2016. Trawling disturbance on megabenthos and sediment in the Barents Sea: chronic effects on density, diversity, and composition. ICES Journal of Marine Science, 73, 98-114. DOI https://doi.org/10.1093/icesjms/fsv200

  5. Cárdenas, P., Rapp, H.T., Klitgaard, A.B., Best, M., Thollesson, M. & Tendal, O.S., 2013. Taxonomy, biogeography and DNA barcodes of Geodia species (Porifera, Demospongiae, Tetractinellida) in the Atlantic boreo-arctic region. Zoological Journal of the Linnean Society, 169 (2), 251-311. DOI https://doi.org/10.1111/zoj.12056
  6. Cebrian, E. & Uriz, M.J., 2007. Contrasting effects of heavy metals and hydrocarbons on larval settlement and juvenile survival in sponges. Aquatic Toxicology, 81, 137-143. 
  7. Cebrian, E., Martı́, R., Uriz, J.M. & Turon, X., 2003. Sublethal effects of contamination on the Mediterranean sponge Crambe crambe: metal accumulation and biological responses. Marine Pollution Bulletin, 46 (10), 1273-1284. DOI https://doi.org/10.1016/S0025-326X(03)00190-5
  8. Chapron, L., Peru, E., Engler, A., Ghiglione, J.F., Meistertzheim, A.L., Pruski, A.M., Purser, A., Vétion, G., Galand, P.E. & Lartaud, F., 2018. Macro- and microplastics affect cold-water corals growth, feeding and behaviour. Scientific reports, 8 (1), 15299-15299. DOI https://doi.org/10.1038/s41598-018-33683-6
  9. Christiansen, S.,  2010. Background document for deep-sea sponge aggregations. OSPAR Commision, Biodiversity Series, 47 pp. https://www.ospar.org/work-areas/bdc
  10. Courtene-Jones, W., Quinn, B., Ewins, C., Gary, S.F. & Narayanaswamy, B.E., 2019. Consistent microplastic ingestion by deep-sea invertebrates over the last four decades (1976–2015), a study from the North East Atlantic. Environmental Pollution, 244, 503-512. DOI https://doi.org/10.1016/j.envpol.2018.10.090
  11. Courtene-Jones, W., Quinn, B., Gary, S.F., Mogg, A.O.M. & Narayanaswamy, B.E., 2017. Microplastic pollution identified in deep-sea water and ingested by benthic invertebrates in the Rockall Trough, North Atlantic Ocean. Environmental Pollution, 231, 271-280. DOI https://doi.org/10.1016/j.envpol.2017.08.026
  12. Edge, K.J., Johnston, E.L., Dafforn, K.A., Simpson, S.L., Kutti, T. & Bannister, R.J., 2016. Sub-lethal effects of water-based drilling muds on the deep-water sponge Geodia barretti. Environmental Pollution, 212, 525-534. DOI https://doi.org/10.1016/j.envpol.2016.02.047
  13. Fallon, S.J., James, K., Norman, R., Kelly, M. & Ellwood, M.J., 2010. A simple radiocarbon dating method for determining the age and growth rate of deep-sea sponges. Nuclear Instruments and Methods in Physics Research Section B: Beam Interactions with Materials and Atoms, 268 (7), 1241-1243. DOI https://doi.org/10.1016/j.nimb.2009.10.143
  14. Fang, J.K.H., Rooks, C.A., Krogness, C.M., Kutti, T., Hoffmann, F. & Bannister, R.J., 2018. Impact of particulate sediment, bentonite and barite (oil-drilling waste) on net fluxes of oxygen and nitrogen in Arctic-boreal sponges. Environmental Pollution, 238, 948-958. DOI https://doi.org/10.1016/j.envpol.2017.11.092
  15. Freese, J.L., 2001. Trawl-induced damage to sponges observed from a research submersible. Marine Fisheries Review, 63 (3), 7-13.

  16. Genin, A., Paull, C.K. & Dillon, W.P., 1992. Anomalous abundances of deep-sea fauna on a rocky bottom exposed to strong currents. Deep Sea Research Part A. Oceanographic Research Papers, 39 (2), 293-302. DOI https://doi.org/10.1016/0198-0149(92)90110-F
  17. Guihen, D., White, M. & Lundälv, T., 2012. Temperature shocks and ecological implications at a cold-water coral reef. Marine Biodiversity Records, 5, e68. DOI https://doi.org/10.1017/S1755267212000413

  18. Henry, L.A. & Roberts, J.M., 2014. Applying the OSPAR habitat definition of deep-sea sponge aggregations to verify suspected records of the habitat in UK waters. JNCC Report No. 508, Joint Nature Conservation Committee, Peterborough, 45 pp. Available from http://data.jncc.gov.uk/data/788783fe-0c27-4faf-9fc3-6b1b1ad3bf78/JNCC-Report-508-FINAL-WEB.pdf
  19. Hoffmann, F., Røy, H., Bayer, K., Hentschel, U., Pfannkuchen, M., Brümmer, F. & de Beer, D., 2008. Oxygen dynamics and transport in the Mediterranean sponge Aplysina aerophoba. Marine Biology, 153 (6), 1257-1264. DOI https://doi.org/10.1007/s00227-008-0905-3
  20. Hoffmann, F., Rapp, H.T., Zöller, T. & Reitner, J., 2003. Growth and regeneration in cultivated fragments of the boreal deep water sponge Geodia barretti Bowerbank, 1858 (Geodiidae, Tetractinellida, Demospongiae). Journal of Biotechnology, 100 (2), 109-118. DOI https://doi.org/10.1016/S0168-1656(02)00258-4
  21. Hogg, M.M., Tendal, O.S., Conway, K.W., Pomponi, S.A., van Soest, R.W.M., Gutt, J., Krautter, M. & Roberts, J.M., 2010.  Deep-sea Sponge Grounds: Reservoirs of Biodiversity. UNEP-WCMC Biodiversity Series No. 32. UNEP-WCMC, Cambridge, UK. https://resources.unep-wcmc.org/products/WCMC_RT401

  22. Howell, K.-L., Piechaud, N., Downie, A.-L. & Kenny, A., 2016. The distribution of deep-sea sponge aggregations in the North Atlantic and implications for their effective spatial management. Deep Sea Research Part I: Oceanographic Research Papers, 115, 309-320. DOI https://doi.org/10.1016/j.dsr.2016.07.005
  23. Isnansetyo, Trijoko, A.,  Setyowati, E.P. & Anshory, H.H., 2009. In vitro Antibacterial Activity of Methanol Extract of A Sponge, Geodia sp. Against Oxytetracycline-Resistant Vibrio harveyi and its Toxicity. Journal of Biological Sciences, 9, 224-230. DOI https://doi.org/10.3923/jbs.2009.224.230
  24. Jørgensen, L.L., Planque, B., Thangstad, T.H. & Certain, G., 2016. Vulnerability of megabenthic species to trawling in the Barents Sea. ICES Journal of Marine Science, 73 (suppl_1), i84-i97. DOI https://doi.org/10.1093/icesjms/fsv107

  25. JNCC (Joint Nature Conservation Committee), 2022.  The Marine Habitat Classification for Britain and Ireland Version 22.04. [Date accessed]. Available from: https://mhc.jncc.gov.uk/

  26. Jones, D.O.B., Gates, A.R. & Lausen, B., 2012b. Recovery of deep-water megafaunal assemblages from hydrocarbon drilling disturbance in the Faroe−Shetland Channel. Marine Ecology Progress Series, 461, 71-82. DOI https://doi.org/10.3354/meps09827

  27. Jones, D.O.B., Hudson, I.R. & Bett, B.J., 2006. Effects of physical disturbance on the cold-water megafaunal communities of the Faroe–Shetland Channel. Marine Ecology Progress Series, 319, 43-54.

  28. Kazanidis, G., Vad, J., Henry, L.-A., Neat, F., Berx, B., Georgoulas, K. & Roberts, J.M., 2019. Distribution of Deep-Sea Sponge Aggregations in an Area of Multisectoral Activities and Changing Oceanic Conditions. Frontiers in Marine Science, 6 (163). DOI https://doi.org/10.3389/fmars.2019.00163
  29. Kędra, M., Renaud, P.E. & Andrade, H., 2017. Epibenthic diversity and productivity on a heavily trawled Barents Sea bank (Tromsøflaket). Oceanologia, 59 (2), 93-101. DOI https://doi.org/10.1016/j.oceano.2016.12.001
  30. Kelmo, F., Bell, J.J. & Attrill, M.J., 2013. Tolerance of Sponge Assemblages to Temperature Anomalies: Resilience and Proliferation of Sponges following the 1997–8 El-Niño Southern Oscillation. PLOS ONE, 8 (10), e76441. DOI https://doi.org/10.1371/journal.pone.0076441
  31. Klitgaard, A.B. & Tendal, O.S., 2004. Distribution and species composition of mass occurrences of large-sized sponges in the northeast Atlantic. Progress in Oceanography, 61 (1), 57-98. DOI http://doi.org/10.1016/j.pocean.2004.06.002
  32. Knudby, A., Kenchington, E. & Murillo, F., 2013. Modeling the distribution of Geodia sponges and sponge grounds in the Northwest Atlantic. PLOS ONE, 8, e82306. DOI https://doi.org/10.1371/journal.pone.0082306
  33. Kutti, T., Bannister, R.J. & Fosså, J.H., 2013. Community structure and ecological function of deep-water sponge grounds in the Traenadypet MPA—Northern Norwegian continental shelf. Continental Shelf Research, 69, 21-30. DOI https://doi.org/10.1016/j.csr.2013.09.011
  34. Kutti, T., Bannister, R.J., Fosså, J.H., Krogness, C.M., Tjensvoll, I. & Søvik, G., 2015. Metabolic responses of the deep-water sponge Geodia barretti to suspended bottom sediment, simulated mine tailings and drill cuttings. Journal of Experimental Marine Biology and Ecology, 473, 64-72. DOI https://doi.org/10.1016/j.jembe.2015.07.017
  35. La Beur, L., Henry, L.-A., Kazanidis, G., Hennige, S., McDonald, A., Shaver, M.P. & Roberts, J.M., 2019. Baseline Assessment of Marine Litter and Microplastic Ingestion by Cold-Water Coral Reef Benthos at the East Mingulay Marine Protected Area (Sea of the Hebrides, Western Scotland). Frontiers in Marine Science, 6 (80). DOI https://doi.org/10.3389/fmars.2019.00080
  36. Lavy, A., Keren, R., Yahel, G. & Ilan, M., 2016. Intermittent Hypoxia and Prolonged Suboxia Measured In situ in a Marine Sponge. Frontiers in Marine Science, 3 (263). DOI https://doi.org/10.3389/fmars.2016.00263
  37. Leys, S.P. & Lauzon, N.R.J., 1998. Hexactinellid sponge ecology: growth rates and seasonality in deep water sponges. Journal of Experimental Marine Biology and Ecology, 230 (1), 111-129. 
  38. Leys, S.P., Kahn, A.S., Fang, J.K.H., Kutti, T. & Bannister, R.J., 2018. Phagocytosis of microbial symbionts balances the carbon and nitrogen budget for the deep-water boreal sponge Geodia barretti. Limnology and Oceanography, 63 (1), 187-202. DOI https://doi.org/10.1002/lno.10623
  39. Leys, S.P., Wilson, K., Holeton, C., Reiswig, H.M., Austin, W.C. & Tunnicliffe, V., 2004. Patterns of glass sponge (Porifera, Hexactinellida) distribution in coastal waters of British Columbia, Canada. Marine Ecology Progress Series, 283, 133-149. DOI http://doi.org/10.3354/meps283133
  40. Luter, H.M., Bannister, R.J., Whalan, S., Kutti, T., Pineda, M.-C. & Webster, N.S., 2017. Microbiome analysis of a disease affecting the deep-sea sponge Geodia barretti. FEMS Microbiology Ecology, 93 (6). DOI https://doi.org/10.1093/femsec/fix074
  41. Malecha, P. & Heifetz, J., 2017. Long-term effects of bottom trawling on large sponges in the Gulf of Alaska. Continental Shelf Research, 150, 18-26. DOI https://doi.org/10.1016/j.csr.2017.09.003

  42. Mariani, S., Uriz, M.-J., Turon, X. & Alcoverro, T., 2006. Dispersal strategies in sponge larvae: integrating the life history of larvae and the hydrologic component. Oecologia, 149 (1), 174-184. DOI https://doi.org/10.1007/s00442-006-0429-9
  43. Mariani, S., Uriz, M.J. & Turon, X., 2003. Methodological bias in the estimations of important meroplanktonic components from near-shore bottoms. Marine Ecology Progress Series, 253, 67-75. DOI https://doi.org/10.3354/meps253067
  44. Masson, D.G., Wynn, R.B. & Bett, B.J., 2004. Sedimentary environment of the Faroe‐Shetland and Faroe Bank Channels, north‐east Atlantic, and the use of bedforms as indicators of bottom current velocity in the deep ocean. Sedimentology, 51 (6), 1207-1241. DOI https://doi.org/10.1111/j.1365-3091.2004.00668.x
  45. McIntyre, F.D., Drewery, J., Eerkes-Medrano, D. & Neat, F.C., 2016. Distribution and diversity of deep-sea sponge grounds on the Rosemary Bank Seamount, NE Atlantic. Marine Biology, 163 (6), 143. DOI https://doi.org/10.1007/s00227-016-2913-z
  46. Morrow, C. & Cárdenas, P., 2015. Proposal for a revised classification of the Demospongiae (Porifera). Frontiers in Zoology, 12 (1), 7. DOI https://doi.org/10.1186/s12983-015-0099-8
  47. Murillo, F.J., Muñoz, P.D., Cristobo, J., Ríos, P., González, C., Kenchington, E. & Serrano, A., 2012. Deep-sea sponge grounds of the Flemish Cap, Flemish Pass and the Grand Banks of Newfoundland (Northwest Atlantic Ocean): Distribution and species composition. Marine Biology Research, 8 (9), 842-854. DOI https://doi.org/10.1080/17451000.2012.682583
  48. Neff, J.M., 2008. Estimation of bioavailability of metals from drilling mud barite. Integrated Environmental Assessment and Management, 4 (2), 184-193. DOI https://doi.org/10.1897/IEAM_2007-037.1
  49. Neff, J.M., 2010. Fate and effects of water based drilling muds and cuttings in cold-water environments. Prepared for Shell Exploration and Production Company, Houston, Texas. Neff & Associates LLC.
  50. Parry, M.E.V., Howell, K.L. , Narayanaswamy, B.E. , Bett, B.J., Jones, D.O.B.,Hughes, D.J., Piechaud, N., Nickell, T.D., Ellwood, H., Askew, N., Jenkins, C. & Manca, E., 2015. A Deep-sea section for the Marine Habitat Classification of Britain and Ireland. JNCC report 530. ISSN 0963 8901 In: JNCC (2015) The Marine Habitat Classification for Britain and Ireland Version 15.03. [Date accessed]. Available from: https://mhc.jncc.gov.uk/

  51. Petralia, R.S., Mattson, M.P. & Yao, P.J., 2014. Aging and longevity in the simplest animals and the quest for immortality. Ageing Research Reviews, 16, 66-82. DOI https://doi.org/10.1016/j.arr.2014.05.003
  52. Pineda, M.-C., Strehlow, B., Sternel, M., Duckworth, A., Haan, J.d., Jones, R. & Webster, N.S., 2017b. Effects of sediment smothering on the sponge holobiont with implications for dredging management. Scientific Reports, 7 (1), 5156. https://doi.org/10.1038/s41598-017-05243-x

  53. Pita, L., Rix, L., Slaby, B.M., Franke, A. & Hentschel, U., 2018. The sponge holobiont in a changing ocean: from microbes to ecosystems. Microbiome, 6 (1), 46. DOI https://doi.org/10.1186/s40168-018-0428-1
  54. Rice, A.L., Thurston, M.H. & New, A.L., 1990. Dense aggregations of a hexactinellid sponge, Pheronema carpenteri, in the Porcupine Seabight (northeast Atlantic Ocean), and possible causes. Progress in Oceanography, 24 (1), 179-196. DOI https://doi.org/10.1016/0079-6611(90)90029-2
  55. Roberts, E.M., Mienis, F., Rapp, H.T., Hanz, U., Meyer, H.K. & Davies, A.J., 2018. Oceanographic setting and short-timescale environmental variability at an Arctic seamount sponge ground. Deep Sea Research Part I: Oceanographic Research Papers, 138, 98-113. DOI https://doi.org/10.1016/j.dsr.2018.06.007
  56. Robertson, L.M., Hamel, J.-F. & Mercier, A., 2017. Feeding in deep-sea demosponges: Influence of abiotic and biotic factors. Deep Sea Research Part I: Oceanographic Research Papers, 127, 49-56. DOI https://doi.org/10.1016/j.dsr.2017.07.006
  57. Rooper, C.N., Wilborn, R., Goddard, P., Williams, K., Towler, R. & Hoff, G.R., 2018. Validation of deep-sea coral and sponge distribution models in the Aleutian Islands, Alaska. ICES Journal of Marine Science, 75 (1), 199-209. DOI https://doi.org/10.1093/icesjms/fsx087
  58. Rooper, C.N., Wilkins, M.E., Rose, C.S. & Coon, C., 2011. Modeling the impacts of bottom trawling and the subsequent recovery rates of sponges and corals in the Aleutian Islands, Alaska. Continental Shelf Research, 31 (17), 1827-1834. DOI https://doi.org/10.1016/j.csr.2011.08.003
  59. Saby, E., Justesen, J., Kelve, M. & Uriz, M.J., 2009. In vitro effects of metal pollution on Mediterranean sponges: Species-specific inhibition of 2′,5′-oligoadenylate synthetase. Aquatic Toxicology, 94 (3), 204-210. DOI https://doi.org/10.1016/j.aquatox.2009.07.002
  60. Scalera Liaci, L. & Sciscioli, M., 1970. The sexual cycle of Erylus discophorus (Schmidt) (Porifera, Tetractinellida). Rivista Di Biologia, 63 (2), 255-270. 
  61. Scanes, E., Kutti, T., Fang, J.K.H., Johnston, E.L., Ross, P.M. & Bannister, R.J., 2018. Mine Waste and Acute Warming Induce Energetic Stress in the Deep-Sea Sponge Geodia atlantica and Coral Primnoa resedeaformis; Results From a Mesocosm Study. Frontiers in Marine Science, 5 (129). DOI https://doi.org/10.3389/fmars.2018.00129

  62. Schönberg, C.H.L., 2016. Effects of dredging on filter feeder communities, with a focus on sponges. Western Australian Marine Science Institution (WASMI) Dredging Science Node Report of Theme 6 – Project 6.1, Western Australian Marine Science Institution, Crawley, WA, 139 pp. Available from https://wamsi.org.au/news/effects-of-dredging-on-filter-feeder-communities-with-a-focus-on-sponges/

  63. Serpetti, N., Narayanaswamy, B.E. & Hughes, D.J., 2014. Assessing the sensitivity of deep-sea sedimentary habitats to pressures associated with marine activities. Phase 2: Review of information on resistance/resilience of key species of ecological groupings identified in Phase 1. JNCC, Peterborough,  [Unpublished report], pp. XX
  64. Sjögren, M., Jonsson, P.R., Dahlström, M., Lundälv, T., Burman, R., Göransson, U. & Bohlin, L., 2011. Two Brominated Cyclic Dipeptides Released by the Coldwater Marine Sponge Geodia barretti Act in Synergy As Chemical Defense. Journal of Natural Products, 74 (3), 449-454. DOI https://doi.org/10.1021/np1008812
  65. Spetland, F., Rapp, H., Hoffmann, F. & Tendal, O., 2007. Sexual reproduction of Geodia barretti (Bowerbank, 1858)(Porifera, Astrophorida) in two Scandinavian fjords. In Custódio, M., et al. (eds.). Porifera Research: Biodiversity, Innovation, Sustainability. Rio de Janeiro: Museu Nacional, pp. 613-620. 
  66. Stévenne, C., 2018. The response of a boreal deep-sea sponge holobiont to an acute crude oil exposure: a mesocosm experiment. Masters thesis, Université de Liège & Havforskningsinstitutet. 
  67. Strand, R., Whalan, S., Webster, N.S., Kutti, T., Fang, J.K.H., Luter, H.M. & Bannister, R.J., 2017. The response of a boreal deep-sea sponge holobiont to acute thermal stress. Scientific reports, 7 (1), 1660. DOI https://doi.org/10.1038/s41598-017-01091-x
  68. Teixidó, N., Garrabou, J., Gutt, J. & Arntz, W.E., 2004. Recovery in Antarctic benthos after iceberg disturbance: trends in benthic composition, abundance and growth forms. Marine Ecology Progress Series, 278, 1-16. DOI https://doi.org/10.3354/meps278001
  69. Tjensvoll, I., Kutti, T., Fosså, J.H. & Bannister, R., 2013. Rapid respiratory responses of the deep-water sponge Geodia barretti exposed to suspended sediments. Aquatic Biology, 19, 65-73.

  70. Trannum, H. C., Nilsson, H.C., Schaanning, M.T. & Øxnevad, S.,  2010. Effects of sedimentation from water-based drill cuttings and natural sediment on benthic macrofaunal community structure and ecosystem processes. Journal of Experimental Marine Biology and Ecology 383 (2), 111-121.

  71. Uriz, M.J., Turon, X. & Becerro, M.A., 2003. Silica Deposition in Demosponges. In Müller, W.E.G. (ed.) Silicon Biomineralization: Biology — Biochemistry — Molecular Biology — Biotechnology, Berlin, Heidelberg: Springer Berlin Heidelberg, pp. 163-193. [Progress in Molecular and Subcellular Biology].
  72. Vad, J., Henry, T.B. & Roberts, J.M., 2017b. Effect of hydrocarbons and dispersants on model shallow-water sponges Halichondria panicea. MASTS Report, Marine Alliance for Science and Technology for Scotland, St Andrews, 8 pp. Available from https://www.masts.ac.uk/media/36455/reportogsg3.pdf
  73. White, M., 2003. Comparison of near seabed currents at two locations in the Porcupine Sea Bight - Implications for benthic fauna. Journal of the Marine Biological Association of the United Kingdom, 83 (4), 683-686. DOI http://doi.org/10.1017/S0025315403007641h

Citation

This review can be cited as:

Last, E.K., Ferguson, M., Serpetti, N., Narayanaswamy, B.E., Hughes, D.J. 2020. Geodia and other massive sponges on Atlanto-Arctic upper bathyal mixed sediment. In Tyler-Walters H. Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 10-10-2024]. Available from: https://www.marlin.ac.uk/habitat/detail/1199

Last Updated: 09/01/2020

  1. Geodia
  2. deep-sea
  3. sponge