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giant barrel sponge environmental impact

Emily McGrath designed the study, performed the field work and analyses, and drafted the manuscript. PERMANOVA with pairwise comparisons for (A) Little Cayman and (B) Lee Stocking Island. By submitting a comment you agree to abide by our Terms and Community Guidelines. Ecol. B. C. & Buss, L. Alleopathy and spatial competition among coral reef invertebrates. Wulff, J. Sponge predators may determine differences in sponge fauna between two sets of mangrove cays, Belize barrier reef. Employing multi-model averaging is widely considered superior in lieu of a priori model choice, but it is possible that our data support another model that we did not examine. Lett. Ecol. Barrel sponges (Xestospongia spp.) To obtain Benzekry, S. et al. 29, 167–177 (2008). Table S1. Thank you for visiting 477, 251–263 (2000). ADS  Cite this article, An Author Correction to this article was published on 27 November 2019. Key Words: Barrel sponge, Xestospongia muta, recovery, reattachment, restoration Introduction The giant barrel sponge, Xestospongia muta, is an abundant, large, and important component of the southeast Florida reef system. Digital images for stereo photogrammetric analysis were taken with a Fujifilm FinePix Real 3D W3 Digital Camera with corresponding underwater camera housing. Models were discarded from further analysis if they failed in predicting reliable asymptotic lengths, or those representative of realistic size estimates based on known sponge sizes39. These studies have all detected shifts in the normally stable microbial communities due to altered environmental conditions and these shifts have simultaneously correlated with a decline in sponge health. Sponges are potentially very long-lived (McMurray et al. Additionally, inorganic nutrient levels steadily increased with depth at LSI but not at LC. 3). Rev. Unlimited viewing of the article/chapter PDF and any associated supplements and figures. Contr. ADS  Assoc. However, we did find a number of barrel sponge recruits during our study challenging this suggestion, although supply was sporadic. Oceanogr. The finger sponge is a sponge with finger-shaped lobes, or branches. Using the model averaged size-at-age curve from the combined Hoga Island sites, the age of the largest sponge measured was estimated to be 27 years old. Barrel sponges (Xestospongia spp.) This research was funded by a Victoria University of Wellington Doctoral Scholarship awarded to Emily McGrath. However, there was no influence of site on specific growth rates or between growth curves, suggesting that despite differences in habitat quality Indo-Pacific Xestospongia spp. The wreck rests on a sand slope adjacent to a shallow reef (8–10 m) and runs from 5 m to 22 m at its base. Effects of the environment on fish juvenile growth in West African stressful estuaries. McGrath, E.C., Woods, L., Jompa, J. et al. Growth and longevity in giant barrel sponges: Redwoods of the reef or Pines in the Indo-Pacific? Ecol. While effort was undertaken to survey a large area, sponges were still chosen haphazardly and it is therefore unlikely that they were the largest and therefore oldest individuals on the wreck. 375, 113–124 (2009). AICc values for each growth model were pooled across sites. 19, 716–723 (1974). This variability, in conjunction with the temporal difference between sponge surveys (2 years herein as compared to 4.5 in the Caribbean)22, may render SGR a less appropriate and robust method of growth estimation in this study. McMurray, S. E., Johnson, Z. I., Hunt, D. E., Pawlik, J. R. & Finelli, C. M. Selective feeding by the giant barrel sponge enhances foraging efficiency. is necessary to confirm their dependence on photosynthetic symbionts and ability to switch to heterotrophy when light availability is low, as well as how this translates into different growth rates. In this instance these analyses, coupled with model selection and MMI based on information theory approach, may reflect higher confidence in growth models over specific growth rate. Clifton, J., Unsworth, R. & Smith, D.) 18–32 (Nova Publishers, 2010). PLoS Comput Biol 10, e1003800 (2014). Seasonality in X. muta growth was reported in the Caribbean22 but was not measured in our study; the volume gained per year in the summer months was in line with growth measured herein from 2014 to 2015 (4,195.53 ± 4,08021; 4,572.60 ± 1,394.66 cm3, Buoy 1). PubMed  1). J. Exp. Further research on the feeding biology of Xestospongia spp. The cube root estimates of sponge volume data from 2014 and 2016 were fitted to the difference equations of the growth models, with the model-averaged result revealing a nearly linear relationship for all sites combined (Supplementary Fig. Despite the ecological importance of barrel sponges there has only been one previous study quantifying their growth rate in the Caribbean; there are currently no estimates of Xestospongia spp. & Dulvy, N. K. Avoiding fishy growth curves. While 5.9% of sponges in this study had a negative growth rate, Caribbean sponges exhibited only positive growth. The impacts of predation and habitat degradation on coral reef sponges. & Zabala, M. Growth dynamics in four Mediterranean demosponges. James Bell co-designed the study and co-authored the manuscript. Biol. A giant barrel sponge living in the backyard of the underwater research vessel Aquarius may be one of the planet's oldest living creatures. Ecol. There was a significant negative correlation with initial sponge volume and specific growth rate (rs = −0.476, P < 0.01). J. Mar. Akaike differences were examined to select models of best fit and these models were used to estimate size-at-age, providing important insight into growth dynamics and potential resilience to environmental perturbations. 357, 139–151 (2008). 18, 303–314 (2015). Henry, L. & Hart, M. Regeneration from injury and resource allocation in sponges and corals–a review. Hydrobiologia 687, 219–226 (2012). Sponges were collected along a depth gradient at Little Cayman (LC) and Lee Stocking Island (LSI), and the microbiome of these samples was analysed using 16S rRNA amplicon sequencing. Model averaged size-at-age for all models combined. López-Legentil S, Song B, Mcmurray SE, Pawlik JR. Bleaching and stress in coral reef ecosystems: Hsp70 expression by the giant barrel sponge Xestospongia muta., DOI: PLoS One 9.1 (2014). The total number of sponges per site used for volumetric analyses were as follows: Buoy 1 (35), Kaldeupa Double Spur (14), Ridge 1 (16), and Sampela 1 (56). Ecosyst. Hydrobiol. Growth and longevity in giant barrel sponges: Redwoods of the reef or Pines in the Indo-Pacific?. Coast. 143, 415–426 (2003). Rev. Surprisingly, Xestospongia spp. It appears that environmental factors that change with depth may contribute to differences in X. muta microbial assemblages, demonstrating the importance of contemporaneous environmental sampling in studies of the microbiome of sponges. Significant pairwise comparisons were not detected at Lee Stocking Island and are not reported. Any queries (other than missing content) should be directed to the corresponding author for the article. PubMed Central  Google Scholar. populations. Previous research on barrel sponges suggests that they should be susceptible to environmental disturbance. 2008;17:1840–9. Mean volume gain was only used for the 2015–2016 sampling event due to the small sample size of Ridge 1 sponges between 2014 and 2015. 1). 50, 799–809 (2005). Estuar. Environmental drivers of microbial community shifts in the giant barrel sponge, Xestospongia muta, over a shallow to mesophotic depth gradient. 84, 175–194 (2006). Data were collected from June to August in 2014, 2015 and 2016. Biol. Although the specific values of the parameters estimated by the model fit may therefore be of moderate confidence, their descriptive power can remain unaffected56. 339, 120–133 (2006). In contrast, using the age at size relationships Caribbean Xestospongia muta would age the biggest sponge on the wreck at approximately 100 years old, much older than the wreck itself (though caution should be taken in such a direct comparison due to the inherent error associated with X. muta age extrapolation)22. $$SGR=({V}_{t}-{V}_{i})\,\ast \,{V}_{i}^{-1}/t$$. López-Victoria, M., Zea, S. & Weil, E. Competition for space between encrusting excavating Caribbean sponges and other coral reef organisms. Automat. Webster, N. S., Cobb, R. E. & Negri, A. P. Temperature thresholds for bacterial symbiosis with a sponge. PubMed  Elephants born in the high stress season have faster reproductive ageing. PubMed Central  Guy Harvey Oceanographic Center Facility. B. density and mean volume was high at a site previously characterized by comparatively higher levels of turbidity and decreased light availability (Supplementary Table 2). Demogr. The authors declare no competing interests. Adviser: Russell Hill, Institute of Marine and Environmental Technology. may be among largest remaining benthic invertebrates on reef systems where coral cover is declining. Efficiently measuring complex sessile epibenthic organisms using a novel photogrammetric technique. These features would be expected to render X. muta susceptible to environmental perturbations. Bio. The oldest giant barrel sponge found off the coast of Venezuela and estimated to be 2300 years old died from SOB in only a few weeks. There was evidence of substantial model support throughout multiple candidate models based on AICc scores such that estimates were strongly supported32. Ecol. Annu. Estuar. Get the most important science stories of the day, free in your inbox. PubMed Google Scholar. Growth curves, however, allow for the intrinsic variability of inter-individual differences in size36. 2. An amendment to this paper has been published and can be accessed via a link at the top of the paper. Comp. in the Indo-Pacific result in a more appropriate comparison to Pines. PubMed  CAS  Mol. 2). From 2015–2016, however, the volume gained at Kaledupa Double Spur was over eight-fold larger than the volumes reported in the Caribbean22 (40,676.99 ± 12,479.05 cm3), though highly variable. However, recent genetic work in this region has revealed a potential cryptic species complex26,27; as such “Xestospongia spp.” will be used throughout. 24, 699–711 (2011). Functional Ecology 31, 2188–2198 (2017). Setiawan, E., De Voogd, N. J., Hooper, J. N. A., Woerheide, G. & Erpenbeck, D. Bottomless barrel-sponge species in the Indo-Pacific? Mar. Several different growth models were identified across sites as supported by the Akaike Information Criterion with a correction for sample size (AICc).There was not one clear model of best fit (wi > 0.9) as the model with the most support, the specialized von Bertalanffy, only had an Akaike weight of 0.533 (Table 1). A recent study found barrel sponges from the Wakatobi Marine National Park to be resilient to the effects of sediment, particularly through the production of mucous, which aids the removal of sediment from the sponge surface61. If you have previously obtained access with your personal account, please log in. Between 2000 and 2012, the giant barrel sponge, Xestospongia muta — which can grow to over a meter tall and wide — covered increasing territory on two reefs off Florida’s Key Largo.The number of sponges per square meter increased on both reefs, on one by an average of 122 percent, researchers report in an upcoming Journal of Experimental Marine Biology and Ecology. Aquat. Prog. Please check your email for instructions on resetting your password. Aquat. Limnol. 81, 229–235 (2006). Front. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate. In order to retrospectively extrapolate Xestospongia spp. In 2014 stereo photogrammetry (as described in the Methods above) was used to measure the volume of 10 haphazardly distributed sponges in an effort to provide an earliest possible date (1963) when Xestospongia spp. In this study, the relationship between the metabolome and the geographical location of sponges within the genus Xestospongia spp. Aiptasia pallida. Pumping rates of the giant barrel sponge, Xestospongia muta on Caribbean reefs: size scaling, environmental controls, and bleaching effects Sponges are among the dominant organisms on coral reefs in terms of diversity, numerical abundance, and biomass. Category. Natl. This study highlights how changes in environmental conditions, such as through degradation, may influence these functionally important species. These sponges also serve as a habitat for many other species such as other invertebrates, benthic fish, bacteria, and … Science (80-.). Despite this highly efficient feeding, it has been suggested that Caribbean X. muta growth is variable and they could live to be hundreds or possibly even thousands of years old22. and you may need to create a new Wiley Online Library account. The HFA Implementation Plan includes programs for: Oyster Restoration, Wetlands, Living Shorelines, Abdul Haris and Jamaluddin Jompa assisted with initial study design, facilitated research permits for work in Indonesia, and final editing of the manuscript. McMurray, S. E., Pawlik, J. R. & Finelli, C. M. Demography alters carbon flux for a dominant benthic suspension feeder, the giant barrel sponge, on Conch Reef, Florida Keys. In the meantime, to ensure continued support, we are displaying the site without styles It is important to note, however, that the picoplankton ingested by sponges29 are likely to have a variety of chlorophyll types, and that dissolved organic matter and heterotrophic picoplankton may provide alternative sources of food. Xestospongia spp. (A) Similarity percentage analysis (SIMPER) for two most significant operational taxonomic units (OTUs) driving differences between the depth range pairs at Little Cayman. Ser. Although there was no direct cause visible in the photographs, the nature of the injuries were suggestive of anchor damage. Abdo, D. A. et al. However, barrel sponges also have the potential to dominate in environments where there is low coral abundance. Shelf Sci. (Springer Science & Business Media, 2003). Variability in sponge growth rates is well reported in the literature; some growth is seasonal21,40,41, and in some species growth measurements are confounded by shrinkage40,41,42. Sponge density was calculated by dividing the total sponge number at each site by the total area sampled during timed roving diver swims of transects approximately 50 m wide (as quantified by pre-measured fin kicks): Buoy 1 (4,500 m2), Sampela 1 (5,450 m2), Kaledupa Double Spur (5,952 m2), Ridge 1 (5,238 m2), Wanci (3,500 m2), Tomea (11,880 m2), Karang Gurita (22,780 m2). Rep. 5, 13946 (2015). Significant effects were investigated further with Tukey post-hoc tests. As such, Xestospongia spp. However, we do not currently have long-term population data on barrel sponges in this region, which would be needed to test this hypothesis. Only 5.9% of sponges showed negative SGRs, the remaining were positive. McMellor, S. & Smith, D. In Marine Research and Conservation in the Coral Triangle: the Wakatobi Marine National Park. Siliceous sponges as a silicon sink: an overlooked aspect of benthopelagic coupling in the marine silicon cycle. Microbiol. Chadwick, N. E. & Morrow, K. M. In Coral Reefs: an ecosystem in transition 347–371 (Springer, 2011). Article  Asterisks (*) and letters denote significant differences between sites. Oceanogr. Mar. It seems reasonable that sponges would not have recruited immediately to the wreck until suitable biofilms had formed, and therefore the age estimates from the wreck provide strong independent support for our size-at-age estimates in the Wakatobi Marine National Park. 23, 1–13 (2014). 4, 353–360 (2013). 72, 5160–5163 (1975). Highly efficient particle retention, coupled with the ability to pump large quantities of water relative to their size, results in the potential to strongly modify water column characteristics by removing a large portion of available particulate food11 and dissolved organic carbon12. can shift to heterotrophic feeding in conditions less favourable to photosynthesis by its symbionts. Publisher's note: Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Next Generation: Daniela Tizabi . Statistically significant shifts in community structure and dissimilarity (∼ 40%) were detected from 10 to 90 m in LC sponges, but a similar shift was not identified in sponges from 10 to 60 m at LSI (only 17% dissimilar). Ecol. Some parameter estimates in the remaining models had moderately large standard errors, likely due to the large parameter number in candidate models (given the apparent linear trend between 2014 and 2016 sizes, Supplementary Fig. Zero long-term net growth has been reported for large, healthy Crambe crambe individuals24. Shelf Sci. In some areas of the Indo-Pacific Xestospongia spp. Can. Xestospongia muta is … Article  & Van Soest, R. W. M. Systema Porifera. Article  At Sampela 1, for instance, competition with other benthic taxa is reduced than at the other sites as the coral cover is low, which would be expected to influence density, mean volume, and potentially growth rates. Mar. 2008) and have important ecological roles in reef ecosystems To view a copy of this license, visit Evol. Bio. The Richards equation produced values identical to those in the generalized von Bertalanffy model and as such was removed to avoid model redundancy36. 222, 1847–1853 (2011). Ecol. The potential intra-specific variation in sponge growth may have the potential to influence parameters such as volume gained and specific growth rates, although this has not been examined in Xestospongia spp. As sponges lack features comparable to otoliths and absolute size-at-age is difficult to quantify in slow growing species, MMI is expected to be particularly useful for reducing model selection uncertainty when estimating sponge growth. 496 (2017). The importance of competitor identity, morphology and ranking methodology to outcomes in interference competition between sponges. Maldonado, M. The ecology of the sponge larva. The USAT Liberty was torpedoed in 1942 during WWII and ran onto shore (8°16′28.48″S; 115°35′35.02″E), where it rested until it sank in 1963 after the tremors associated with the eruption of Mount Agung caused the ship to slip into the sea. SGR decreased with increasing size. A distinction of note between this practice and the data presented herein, however, is the ability to “ground truth” using actual size-at-age data, typically otoliths and fish length data39,55. 9, 178–187 (2008). All significantly contributing OTUs are reported for each pair. 3). Understanding barrel sponge demography is particularly important given current trends of habitat degradation in the Indo-Pacific and elsewhere. Article  A guide to the classification of sponges. Integr. Sponges ranged in volume from approximately 80,000 cm3 to 310,000 cm3 (n = 10). Cleary, D. F. R. & De Voogd, N. J. The cause of SOB is unknown, but evidence suggests that it is a result of a change in environmental factors, particularly rising water temperatures. de Goeij, J. M. et al. & Jamieson, I. G. Multimodel inference in ecology and evolution: challenges and solutions. Hometown: Silver Spring, Maryland. The size of an organism, and the population within which it resides, will likely affect the magnitude of its influence on other organisms3. MATH  & Fialkov, V. A. Trophic effects of sponge feeding within Lake Baikal’s littoral zone. Biol. Lesser, M. P. Benthic – pelagic coupling on coral reefs: Feeding and growth of Caribbean sponges. Interestingly, SGRs were slightly slower than that of X. muta, yet growth models supported rapid growth; published estimates of comparably sized X. muta were over twice as old as Indo-Pacific sponges (53–55 as compared to 23 years of age, respectively), although extrapolation errors are likely to increase with sponge size. Until recently, Indo-Pacific barrel sponges were believed to solely include X. testudinaria, X. exigua, and X. berguista. Fish. Methods Ecol. In particular, the importance of spatial competition in shaping the ecology of sponges has received considerable attention49,50,51,52,53,54, and may have a large influence on the dynamics of Xestospongia spp. Use the link below to share a full-text version of this article with your friends and colleagues. OTUs that composed principal coordinate analysis (PCoA) clusters 13 and 38 from the Little Cayman (LC) dataset. The range of SGRs herein was also larger (−0.12–6.24 yr−1 as compared to 0.02–4.04 yr−1 in the Caribbean)22. Sociobiol. Barthel, D. & Tendal, O. S. The sponge association of the abyssal Norwegian Greenland Sea: species composition, substrate relationships and distribution. Wulff, J. L. Ecological interactions of marine sponges. Based on bulk stable isotopic variability, sponges collected from LC were generally more enriched in 15N and less enriched in 13C as depth increased, suggesting a transition from dependency on photoautotrophy to heterotrophy as depth increased. Mortality estimates were possible only at Buoy 1 and Sampela 1 and were variable; the highest mortality recorded was 11 individuals in one year (Sampela 1, 2014) and the lowest 3 (Buoy 1, 2015). Giant barrel sponges in the genus Xestospongia may be among the largest benthic invertebrates providing habitat and fulfilling ecosystem services on reefs where coral is declining. Understanding the life history traits of an organism including growth, recruitment, and mortality are central to quantifying its contribution to ecosystem functioning1, as well managing species in response to environmental perturbations2. Classical mathematical models for description and prediction of experimental tumor growth. Tube, barrel, and finger sponges have vertical morphologies. For example, individuals of Ianthella basta nearly 2 m high were reported to be only 10 years old58. Google Scholar. Ehrlén, J. PhD Thesis (2013). & Morris, W. F. Predicting changes in the distribution and abundance of species under environmental change. The reduction in coral cover at Sampela 1 is largely thought to have occurred in the last 10–20 years, with coral cover declining from 30% to <8%60, which will likely have released barrel sponges from spatial competition and potentially allowed a larger population size. Oceanogr. Shelf Sci. As both SGRs and growth curves reflect mean growth rates, the conflicting results reported herein highlights the importance of long term monitoring for Indo-Pacific Xestospongia spp., as well as the need for further investigation into the parameters which shape the growth of these ecologically important sponges. 12, 631–639 (1998). Surviving in a marine desert: the sponge loop retains resources within coral reefs. Scientific Reports 52, 948–958 (2007). It is possible that barrel sponges have specific adaptions to live in the sedimented conditions at Sampela 1 that support their success. The giant barrel sponge, Xestospongia muta, is a high microbial abundance sponge found on Caribbean coral reefs along shallow to mesophotic depth gradients where multiple abiotic factors change with depth. Resilience to disturbance despite limited dispersal and self-recruitment in tropical barrel sponges: implications for conservation and management. As we were unable to differentiate which species were surveyed, and due to the demographic nature of the study, all sponges were treated at the level of genus. The mean specific growth rate (SGR) for the Caribbean barrel sponge X. muta was reported as 0.52 ± 0.65 year−1 22 (±SD), as compared to 0.47 ± 0.07 year−1 (±SE) in the present study. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. Limnol. The giant barrel sponge (Xestospongia muta) can live to be 2,300 years old, like the oldest known redwood tree. Molecular analyses are required in order to clarify this issue and fully describe the nature of the species complex. 3). Perea-Blazquez, A., Davy, S. K. & Bell, J. J. 15, 347–400 (2006). Bull. Garrabou, J. Aquat. : specialized von Bertalanffy, generalized von Bertalanffy, Gompertz, Richards, Tanaka (Supplementary Table 2). Volume was calculated by approximating geometric shapes for each sponge shape and corrected for spongocoel volume22 (described in the Supplementary Information). The vase sponge is characterized by a large bell shape with a deep central cavity. They have a large body size, low population connectivity28, slow growth, are likely long-lived22, and have high larval mortality common to broadcast spawning59. Furthermore, the recognized Indo-Pacific Xestospongia species complex comprised of cryptic species26,27 may also confound measurements of growth. Sponges from different sites showed differences in density, volume gained, and mean volume, but not growth rates. Four ‘core’ sites were surveyed around Hoga Island in the Wakatobi Marine National Park (WMNP) between 2014 and 2016: Sampela 1, Ridge 1, Buoy 1, and Kaledupa Double Spur (Fig. in other areas of Sulawesi have been reported to be constrained to deeper depths46, supporting a limited reliance on symbionts. Google Scholar. Spongocoels were categorized as either cylinders or inverted truncated elliptical cones (depending on the sponge) and volume was calculated accordingly. Giant Barrel Sponge (Xestospongia muta) Bonaire ... say, environmental challenges, some sponges can migrate – at an astonishing rate of a few millimeters per day. Int. Acorn barnacles live along rocky shores throughout the north Atlantic and north Pacific oceans. Lisa Woods assisted in analyses. 90, 125–158 (2005). Interestingly, results demonstrate that Indo-Pacific barrel sponges achieve a comparable size to that of their Caribbean cohorts much faster, and therefore large barrel sponges on Indo-Pacific reefs are more comparable to Pine trees rather than the Redwoods proposed in the Caribbean23. Barrel sponge recruitment was variable and ranged across sites and years from no recruits (Kaledupa Double Spur: 2015, 2016 and Ridge 1, 2015) to 20 recruits (Buoy 1, 2016; Supplementary Fig. However, despite these features we found barrel sponges were larger and more abundant at a low quality site (as determined by published levels of high levels of sedimentation and turbidity and low coral cover, Supplementary Table 2). These specimens may be over 100 years old, as the sponges grow only about 1.5 cm a year. IEEE Trans. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. Mar. Abiotic and biotic factors alike are likely to affect the population and individual sponge dynamics at each site; these include factors not measured in this study such as food supply and consumption, hydrodynamics, spatial competition, wide-scale disturbance (e.g. & Barnes, D. K. A. Three further sites were surveyed in 2014 to estimate population densities in the wider WMNP: Karang Gurita, Wanci Harbor, and Tomea (Fig. Growth dynamics and mortality of the encrusting sponge Crambe crambe (Poecilosclerida) in contrasting habitats: correlation with population structure and investment in defence. Estuar. Correspondence to Here the demography of Xestospongia spp. We then used our size-at-age model to estimate the approximate age of the sponges on the wreck and compared these to the known maximum possible sponge age. Meesters, E. H., Hilterman, M., Kardinaal, E., Keetman, M. & Bak, R. P. M. Colony size-frequency distributions of scleractinian coral populations: spatial and interspecific variation. Modell. This makes it the longest living animal. Variation in age extrapolated from growth model projections is not uncommon due to differences in model selection, sampling methods, reproductive life history or morphological differentiation57; the importance of model choice and parameter uncertainties are detailed in the Supplementary Information. However, it is currently unknown whether such characteristics are applicable to Xestospongia spp.

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